Bryophyta by O. P. Sharma

Series on Diversity of Microbes and Cryptogams
Bryophyta
O P Sharma, with his 36 research articles published in national and
international journals, 33 books written for university students, and 40
years of teaching experience, is an able researcher, established Indian
author, and an experienced teacher. His areas of research include pollen
morphology, angiosperm’s anatomy and mycology, with special focus on
Indian Cyperaceae (with particular interest on Cyperus). Over a dozen of
Dr Sharma’s books have been published through internationally known
publishers. He has also revised Economic Botany, an internationally
renowned text by the late Professor Albert F Hill (Harvard University,
USA), a publication of McGraw-Hill, New York.
Encouraging reviews of his books have been published in reputed
scientific journals such as Current Science, and his books on Practical
Botany have received appreciations from some eminent botanists including J D Dodge (England),
T Christensen (Denmark), J M Herr (USA) and C R Metcalfe (England).
Immediately after completing his postgraduation (MSc) in Botany with first division from CCS
University, Meerut, and thereafter obtaining a PhD from the same university, Dr Sharma started his
teaching career in 1967 as a faculty member of the Botany Department, Meerut College, Meerut,
and retired from active services as a Reader from the same department in 2007. Besides attending
several national and international workshops, symposia and conferences during the four decades of his
teaching career, Dr Sharma is still enjoying his post-retirement innings as an active author.
Recently, Dr O P Sharma revised some of his widely circulated books published through
McGraw Hill Education, which include Plant Taxonomy (released first in 1993 and reprinted 19 times);
Textbook of Algae (released first in 1986 and reprinted 20 times), now titled Algae; and Textbook of
Fungi (released first in 1989 and reprinted 17 times), now titled Fungi and Allied Microbes. His latest
published books through McGraw Hill Education are Pteridophyta (published in 2012) and the present
one titled Bryophyta (being published in 2013).
Series on Diversity of Microbes and Cryptogams
Bryophyta
O P Sharma
Reader (Retired)
Department of Botany
Meerut College, Meerut
McGraw Hill Education (India) Private Limited

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Series on Diversity of Microbes and Cryptogams: BryOPhyta

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Preface xvii
1.1 What are Bryophytes? 1

1.2 How do Recent Systematists Treat the Terms “Bryophyta”, “Bryophytes” and
“Bryobiotina”? 3
1.3 Why are Bryophytes Always Small-Sized and Lack True Leaves? 3
1.4 Why do Bryophytes Possess Rhizoids Instead of Roots? 4
1.5 Number of Species and Gene-Bank Database 4
1.6 Age of Bryophytes 4
1.7 Occurrence and Distribution 5
1.8 Gametophyte (Plant Body) 5
1.9 Vegetative Reproduction 6
1.10 Sexual Reproduction 7
1.11 Sporophyte 9
1.12 Young Gametophyte 10
1.13 Life-Cycle (Alternation of Generations) 10
1.14 A Note on Regeneration in Bryophytes 11
1.15 Salient Features of Bryophytes: At a Glance 11
1.16 Affinities of Bryophytes 12
1.17 Origin of Bryophytes 15
Test Your Understanding 15
2.1 Classification 16
2.2 Major Differences Between Classes of Bryophyta 17
2.3 Latest Position of Classification of Bryophytes 19
3.1 What are Hepaticopsida (= Liverworts)? 22
3.2 Why are They Called Liverworts or Hepaticopsida? 22
3.3 General Characteristics 23
3.4 Classification of Hepaticopsida 23
3.5 Takakiales 24
3.6 Takakia 25
3.7 Calobryales 28
3.8 Calobryum and Haplomitrium 29

4.3 Sub-Order Jungermannineae (Jungermanniales Acrogynae) 35
4.4 Porellaceae 36
4.5 Porella 36
4.6 Frullaniaceae 45
4.7 Frullania 46
4.8 Sub-Order Metzgerineae (Jungermanniales Anacrogynae) 52
4.9 Pelliaceae 52
4.10 Pellia 52
4.11 Riccardiaceae 59
4.12 Riccardia 60
4.13 Fossombroniaceae 65
4.14 Fossombronia 65
5.1 Sphaerocarpales and their General Characteristics 70

5.2 Sphaerocarpos 70
5.3 Riella 74
6.1 What are Monocleales? 77

6.2 General Characteristics of Monocleales 77
6.3 Monoclea 78
7.1 Limits of Marchantiales? 81

7.2 Common Genera 81
7.5 Ricciaceae 83
7.6 Riccia 83
7.7 Marchantia 94
7.8 Comparison of Sporogonium of Riccia and Marchantia 111
7.9 Origin and Evolution of Marchantiaceous Thallus 111
8.1 What is Anthoceropsida? 116

8.4 Anthocerotaceae 117
8.5 Anthoceros 118
8.6 Apospory in Anthoceros 129
8.7 Notothylaceae 130
8.8 Notothylas 131
8.9 Biological Importance of the Sporophyte of Anthoceros 133
8.10 Affinities of Anthocerotales 135
8.11 Difference between Anthocerotopsida and Hepaticopsida 137
8.12 A Note on the Origin of Anthocerotales 138
9.1 What is Bryopsida? 140

9.4 Distribution and Habitat 141
9.5 Habit 142
9.6 Vegetative or Asexual Reproduction 143
9.7 Apospory in Mosses 143
9.8 Sex Organs 144
9.9 Resemblances and Differences between Hepaticopsida (Liverworts)
and Bryopsida (Mosses) 147
10.1 Sphagnales or Bog Mosses 150

10.2 Sphagnum 151
10.3 Affinities of Sphagnum 161
10.4 Andreaeales 162
10.5 Andreaea 163
10.6 Bryales 166
10.7 Funaria 166
10.8 Buxbaumia: A Botanical Note 181
10.9 Tetraphis: A Botanical Note 183
10.10 Archidium: A Botanical Note 185
10.11 Pogonatum: A Botanical Note 186
10.12 Polytrichum 189
11.1 What is a Gametophyte? 197

11.2 External Form of Mature Gametophyte 197
11.3 Thalloid Forms of Gametophyte 198
11.4 Leafy Forms of Gametophyte 198
11.5 Appendages Found on the Gametophyte 200
11.6 Internal Form of Mature Gametophyte 201
12.1 What is a Sporophyte and a Spore? 205

12.2 How is the Sporophyte Different from the Sporogonium? 205
12.3 Bryophytic Sporophyte: A Body Dependent on Gametophyte 206
12.4 Structure of Sporophyte 206
12.5 Development and Differentiation of Different Organs of Sporophyte 208
13.1 What is Technically a Spore? 218

13.2 Are Bryophytes Homosporous or Heterosporous? 218
13.3 Size and Output of Spores 219
13.4 Structure of Spores of Bryophytes 219
13.5 Spore Germination and Formation of Gametophyte in Liverworts (Hepaticopsida) 219
13.6 Spore Germination and Formation of Gametophyte in Hornworts (Anthoceropsida) 222
13.7 Spore Germination and Formation of Gametophyte in Mosses (Bryopsida) 222
13.8 Spore Viability 224
13.9 Polarity in Spore Germination 224
14.1 Major Factors Affecting Sexuality 226

14.2 Influence of Photoperiod on Sexuality 226
14.3 Influence of Temperature on Sexuality 227
14.4 Influence of Carbohydrates on Sexuality 228
14.5 Influence of Nitrogenous Substances on Sexuality 229
14.6 Influence of pH and Hydration of Medium on Sexuality 229
14.7 Influence of Growth Regulators on Sexuality 229
15.1 What is Alternation of Generations? 232

15.2 Who Discovered Alternation of Generations in Archegoniatae? 233
15.3 What are Antithetic Alternation and Homologous Alternation of Generations? 233
15.4 Phenomena of Apogamy and Apospory 233
15.5 Antithetic Theory 234
15.6 Final Picture about Nature and Evolution of Primitive Sporophyte 240
15.7 Homologous Theory 241
15.8 Genetical and Ontogenetic Views on Alternation of Generations 241
16.1 Current Status of Molecular Studies on Bryophyte Phylogeny 243

16.2 Bryophytes and Genosystematics Studies 243
16.3 Macrosystematics and Origin of Bryophytes 244
16.4 Marchantiophyta or Liverworts 245
16.5 Anthocerotopsida or Hornworts 245
16.6 Bryophyta or Mosses 246
16.7 Major Roles of Molecular Phylogenetics 248
17.1 What is Morphogenesis? 250

17.2 Limits of Morphogenetic Studies in Bryophytes 250
17.3 Germination of Spore 250
17.4 Initiation of Bud and its Growth into Gametophyte 252
17.5 Physiology of Sex Expression 252
17.6 Growth of Diploid Sporophyte 253
17.7 Vegetative Propagation 253
17.8 Metabolism 254
17.9 Senescence 255
17.10 Physiology of Rhizoid Formation 255
18.1 Difference between Vegetative Reproduction and Regeneration 257

18.2 Methods of Vegetative Reproduction in Bryophytes 257
18.3 Botanical Note on Regeneration in Bryophytes 261
19.1 What is Archesporium? 264
19.2 Products of Archesporium 264
19.3 Fundamental Embryonic Layers of Capsule 266
19.4 Origin, Position and Fate of Archesporium in Some Selected Genera 266
19.5 A Note on the Origin of Elaters 267
20.1 Apogamy and Apospory: Two Alternative Pathways of Life Cycle 269
20.2 Why are the Phenomena of Apogamy and Apospory of Considerable Interest? 269
20.3 Apogamy 270
20.4 Apospory 273
21.1 Water Relations 275

21.2 Growth Forms in Bryophytes 276
21.3 Water-Holding Capacity of Bryophytes 277
21.4 Photosynthesis 277
21.5 Nitrogen Metabolism 278
21.6 Respiration 278
21.7 Enzymes 279
21.8 Organic Acids 279
21.9 Movements 279
22.1 Antibiotics 281

22.2 Growth Substances 282
22.3 Lipids 282
22.4 Alkanes 282
22.5 Fatty Acids 283
22.6 Terpenoids 283
22.7 Flavonoids 283
22.8 Lignins 283
22.9 Carotenoids 284
22.10 Carbohydrates 284
22.11 Organic Acids 284
22.12 Enzymes 285
22.13 Amino Acids 285
22.14 Allergenic and Anti-Tumour Activities 285
23.1 Indicator Species of Bryophytes 286
23.2 Erosion Control by Bryophytes 287
23.3 Nitrogen Fixation 287
23.4 Bryometer or Moss Bag 288
23.5 SO2 and Acid Rain 288
23.6 Bryophytes as Bioindicators of Heavy Metals in Air Pollution 289
23.7 Copper Mosses as Indicators of Copper Concentration 290
23.8 Bryophytes as Bioindicators of Some Other Air Pollutants 290
23.9 Bryophytes as Indicators of UV-B Radiation 291
23.10 Bryophytes as Indicators of Radioactivity 291
23.11 Bryophytes as Indicators in Aquatic Habitats 291
23.12 Bryophytes as Cleaning Agents of Toxic Waste 292
24.1 What are Biologically Active Compounds? 293

24.2 Some Biologically Active Compounds Extracted from Bryophytes 293
24.3 Characteristics of Biologically Active Compounds Isolated from Bryophytes 294
24.4 Characteristic Scents 295
24.5 Pungency and Bitterness 295
24.6 Dermatitis 296
24.7 Cytotoxic, Anti-HIV-1, and DNA Polymerase B-Inhibitory Activity 296
24.8 Antifungal and Antimicrobial Activity 297
24.9 Insect Antifeedant, Mortality and Nematocidal Activity 297
24.10 Superoxide Anion Radical Release Inhibitory Activity 297
24.11 Enzymes and Nitric Oxide Production Inhibitory Activity 298
24.12 Pesticidal and Plant-Growth Inhibitory Activity 298
24.13 Neurotrophic Activity 298
24.14 Muscle-Relaxing Activity 299
24.15 Inhibitory Activity Against Osteoporosis and Allergy 299
24.16 Cardiotonic and Vasopressin Antagonist Activity 300
24.17 Antiobesity Activity 300
24.18 Synthesis of Some Bioactive Compounds from Constituents of Liverworts 300
25.1 Role of Plants in Regulating Biogeochemical Cycles 302

25.2 Do Bryophytes have the Same Role as Vascular Plants in Regulating
Biogeochemical Cycles? 302
25.3 How do Bryophytes Influence Ecosystem Functions? 303
25.4 Major Roles of Bryophytes in C- and N-Cycling 303
25.5 Ability of Bryophytes to Influence Local Soil Climates 303
25.6 Role of Bryophytes in Carbon Cycling 304
25.7 Role of Bryophytes in Nitrogen Cycling 307
25.8 What More can be Done in Understanding Role of Bryophytes in C and
N Cycling? 308
26.1 Views on the Evolution of the Gametophyte in Bryophytes 310

26.2 Mechanism of Retrogressive Evolution 310
26.3 Theory of Progressive Evolution 312
27.1 What is Evolution of the Sporophyte in Bryophytes? 314

27.2 Theory of Progressive Sterilization of Potentially Sporogenous Tissue 314
27.3 Theory of Progressive Simplification 318
28.1 How do Bryophytes Maintain Poor Internal Conducting Strands? 320

28.2 External Conduction and its Significance 320
28.3 Internal Conduction and its Significance 322
28.4 Anatomy of Midrib and Leaf Traces 325
28.5 Similarities and Differences Between Hydroids of Bryophytes and Tracheary
Elements of Primitive Vascular Plants 326
28.6 Similarities and Differences Between Leptoids of Polytrichales and Sieve
Elements of Vascular Plants 326
28.7 Internal Conduction of Water by Gametophyte 327
28.8 Internal Conduction of Water by Sporophyte 327
28.9 Conduction of Organic Compounds in Bryophytes 327
29.1 Some Basic Techniques of Cytogenetics 329

29.2 Some Studies on Mitosis 330
29.3 Some Studies on Meiosis 330
29.4 Heterochromatin 332
29.5 Chromosome Number 332
29.6 Extra Chromosomes and Sex-Associated Chromosomes 334
29.7 Hybridization 336
29.8 Polyploidy 337
29.9 Cytotaxonomy 338
30.1 Views of Johannes et al. (2007) on Ecology of Bryophytes 340
30.2 Limits of Ecology of Bryophytes 340
30.3 Autecology 341
30.4 Synecology 341
30.5 Succession 343
30.6 Factors of the Habitat 344
31.1 Origin of Bryophytes 346

31.2 A Note on the Monophyletic or Polyphyletic Nature of Bryophytes 346
31.3 Origin of Bryophytes from Pteridophytes 347
31.4 Origin of Bryophytes from Algae 348
31.5 Fossil History of Bryophytes 349
32.1 Utility of Bryophytes: A General Overview 353

32.2 Ecological Uses of Bryophytes 353
32.3 Medicinal Uses of Bryophytes 353
32.4 Bryophytes as Food Source 355
32.5 Horticultural Uses of Bryophytes 356
32.6 Household Uses of Bryophytes 356
32.7 House Construction 356
32.8 Moss Industry 357
32.9 Fuel from Bryophytes 357
32.10 Pesticides from Bryophytes 358
32.11 Clothing from Bryophytes 358
32.12 Bryophytes Used for Culturing 358
32.13 Bryophytes as Seed Beds 359
32.14 Moss Gardens 359
32.15 Bryophytes and Genetic Engineering 359
Bryophyta is the next and last of the Series on Diversity of Microbes and Cryptogams, and my main
aim of writing this book is to fill a gap which exists between the full, straightforward treatment of the
morphology of bryophytes and research literature in all its details and diversity of topics. My hope is
that it will enable the university students to see morphological and other related bryological facts from
a new angle and at the same time have their interest directed to other branches of botany.
Another major aim of writing a book on Bryophyta is evidenced by Appendix-IV of this book
(Globally Published Books on Bryophyta), which mentions a list of approximately 120 books. Out
of this list, only three books are available in the Indian market which suit, to some extent, the syllabi
requirements of UG and PG students of Indian universities, but none of these three has been revised
during the last 10 to 15 years. This, in itself, provides sufficient ground for writing and publishing
a new book on Bryophyta, which discusses and explains the subject matter in the light of recent
developments in bryology. This is not only a need of the students but also their right to study the
subject in the light of recent global developments. The contents of this book were prepared to fulfill
this requirement of the students of Indian universities and of the Indian subcontinent, and the book in
its present form is the result.
Furthermore, Bryophyta is a syllabi requirement of botany students as a compulsory full paper at the
postgraduate level and half of a paper at the undergraduate level in all universities of the country. Due
to this too, a new book of such nature is a ‘must’ and a ‘need of the hour’ for Indian students.
The present book on Bryophyta includes 32 chapters.

The initial two chapters deal with the basics of Bryophyta and a glimpse of their classification,
followed by eight chapters (3 to 10) discussing the major aspects and life-history details of almost all
major genera of all the three bryophytic groups (i.e. Hepaticopsida, Anthoceropsida and Bryopsida).
They are followed by the remaining 22 chapters (11 to 32) discussing almost all important general
topics of Bryophyta.
Four appendices are given at the end of the book. Appendix I provides “answers to all short-answer
questions” of all the chapters of the book. Appendix II compares different aspects of all major and
commonly available bryophytic genera in the form of some comparative tables. Appendix III explains
the meanings/definitions of 233 bryological terms in simple language, and Appendix IV gives a list
of about 120 globally published books and theses on Bryophyta. Details of research publications have
been intentionally avoided, with the feeling that the book should not look too cumbersome to young
students.
Some major highlights of the book are as follows:

Complete coverage to general aspects of all groups of Bryophyta:
Hepaticopsida (including Takakiales, Calobryales, Jungermanniales, Sphaerocarpales,
Monocleales and Marchantiales)
Anthoceropsida
Bryopsida
Complete coverage to all important topics in Bryophyta: Classification, Physiology,
Reproduction, Regeneration, Constituents, Origin and Evolution, Ecology, Economic
Importance
Detailed life-histories of:
13 liverworts, 2 hornworts and 8 mosses
Specific life-history details of over a dozen genera included in the syllabi for postgraduate
students (Takakia, Calobryum, Haplomitrium, Frullania, Riccardia, Fossombronia,
Sphaerocarpos, Riella, Monoclea, Notothylas, Andreaea, Buxbaumia, Tetraphis and
Archidium)
Complete chapter-wise coverage of all important general topics in bryology:
Gametophytes of bryophytes
Sporophytes of bryophytes
Alternation of generations
Vegetative reproduction in bryophytes
Origin and fate of archesporium in bryophytes
Apospory and apogamy in bryophytes
Physiology of bryophytes
Evolution of gametophyte in bryophytes
Evolution of sporophyte in bryophytes
Ecology of bryophytes
Origin of bryophytes and their fossil history
Six independent chapters on new and modern topics, not available in most Indian books
on bryology:
Sexuality in Bryophytes
Phylogeny of Bryophytes: Role of Genosystematics
Chemical Constituents of Bryophytes
Bryophytes as Indicators of Environmental Conditions and Pollution
Biologically Active Compounds from Bryophytes
Conduction in Bryophytes
Five application-based chapters:
Economic Importance of Bryophytes
Morphogenesis
Cytogenetics and Cytotaxonomy of Bryophytes
Role of Bryophytes in Carbon and Nitrogen Cycling
Classification of Bryophytes
rich pedagogy:
Profusely illustrated with more than 200 well-labelled diagrams
Examination preparation tools to help students:
More than 500 chapter-end questions, patterned on university questions, with answers to
select questions in Appendix I
18 comparative tables spread across the text and at the end of the book in Appendix II
Glossary of over 230 bryological terms explained in Appendix III
List of 120 globally published books on Bryophyta in Appendix IV
Pictorial life-cycles of dozens of genera
In view of the above-mentioned highlights, I can confidently say that the present book caters to all
needs of readers, particularly undergraduate and postgraduate students of botany as well as agriculture
of all universities the world over, in general, and of Indian universities, in particular. It should also
prove useful to students preparing for AIPMT, CPMT, IAS, IFS, PCS, NET, SLET and several other
major competitive examinations of the country.
Prior to its publication process, the manuscript was sent for review to many subject experts/reviewers.
They took pains in critically reviewing and examining the manuscript, and for this I express my heartfelt
thanks to all of them.
At this stage, I also wish to express my special thanks to Professor M U Charaya of CCS University,
Meerut, for the desired help in searching and supplying me some valuable literature from the library
and the Internet; my wife, Dr (Mrs) Kanti D Sharma, PhD, for her unfailing assistance and patience;
and my lovely and dearest grandchildren (Kuhu and Karan) for allowing me time that was due only to
them.
Last, but not the least, I wish to put on record my most sincere thanks to the entire editorial and
production teams of McGraw Hill Education (India). Without their support, my this ninth book from
McGraw Hill Education (India) under their Higher Education Programme, would have not seen the
light of day. I express my gratitude to all of them.
Comments and suggestions for the improvement of this book may be sent to the publisher’s email,
given below, or directly to me.
O P SharMa
+91-9837566555
+91-121-2401400
Do you have any further request or a suggestion? We are always open to new ideas (the best ones come
from you!). You may send your comments to tmh.sciencemathsfeedback@gmail.com
1
Introduction
WHAT ARE BRYOPHYTES? 1.1

Bryophytes (Greek—bryon—moss; phyton—plant) include liverworts, hornworts and mosses, and
occupy a position in the plant kingdom in between thallophytes and pteridophytes. These are land-
inhabiting plants, restricted mostly to moist, shady places, and totally dependent on external water for
completing their life-cycle. Because of these characters, bryophytes are also called “amphibians of the
plant kingdom”. The name ‘bryophyta’ was first introduced by Braun(1864). Recently, Crum(2001) in
his Structural Diversity of Bryophytes stated that “ in the 1600’s, Jung considered mosses to be aborted
plant foetuses! Today, they occupy a position within the plant kingdom and may even be considered to
have their own subkingdom”.
Engler (1886) opined that all the plants above the level of Thallophyta (i.e. algae, fungi and lichens)
should be grouped under the subkingdom Embryophyta. The Embryophyta (or Embryobionta) comprises
all those plants in which a multicellular embryo develops from the zygote while the zygote is still
attached to the parent. The Embryophyta includes Bryophyta, Pteridophyta and Spermatophyta. The
Pteridophyta and Spermatophyta (i.e. gymnosperms and angiosperms) possess vascular tissues and are
often grouped together under Tracheophyta. Bryophytes do not possess vascular tissues. Bryophytes
may, therefore, be defined as “the embryophytes that do not possess vascular tissue.”
Bryophytes are photosynthetic, nonvascular plants which show clear heteromorphic alternation of
generations. Gametophytic generation is dominant, well-developed and independent. Gametophytes
are either thalloid or foliose, i.e. bear leaflike structures. The sporophyte is never independent. It is
always dependent on the gametophyte. All bryophytes require water for the process of fertilization.
No vascular tissue is present in either generation. True roots, stems and leaves are absent in either
generation in all bryophytes. Cuticle and stomata are absent in the leaflike structures of mosses.
Bryophytes have also been described variously by different workers as under:
As defined in the 1989 edition of the Chambers Biology Dictionary of Cambridge University
Press, Bryophyta is a “division of the plant kingdom containing about 25,000 species of small,
rootless, thalloid or leafy nonvascular plants; includes the liverworts (Hepaticopsida), the hornworts
(Anthoceropsida) and the mosses (Bryopsida), having alternation of generations in which the
2 � Bryophyta
gametophyte is the dominant generation, the sex organs are archegonia and antheridia and the
sporophyte is more or less parasitic on the gametophyte”.
In the Penguin Dictionary of Botany, Bryophyta has been described as “a division of nonvascular
plants, mainly terrestrial in habitat.” It comprises three classes—Hepaticae (liverworts), Musci
(mosses) and Anthocerotae (hornworts). Bryophyta are generally small, low-growing plants, in most
cases susceptible to desiccation and hence limited to damp or humid environments. Their life-cycle
shows a heteromorphic alternation of generations with the haploid gametophyte, which may be homo-
or heterothallic, the dominant generation. The ephemeral sporophyte is partly or completely parasitic
on gametophyte ... .” Close resemblances between some bryophytes (e.g. mosses) and algae, especially
between moss protonema and algal filaments “suggest that bryophytes evolved from algae, most
probably green algae”, since both of them “share similar photosynthetic pigments, food reserves and
cell-wall constituents”. Bryophytes, however, show more “morphological differentiation than the algae
and differ also in producing aerial spores and having enclosed sex organs”. They also lack roots, and
water is required by them for the dispersal of their male gametes and also for the fertilization process.
Bryophytes depend on water for their process of fertilization. Their ciliate antherozoids have to swim
in water drops to effect fertilization. Due to these characteristics, bryophytes are also called “amphibians
of the plant kingdom.” Recently, Daniel Norris (2003), in an interesting article in Fremontia, defines
bryophytes as “green plants without flowers and fruits, and lacking a well-defined system of vascular
tissue for transporting plant fluids throughout the plant. They reproduce, not by seeds, but by single-
celled spores. Mosses and most liverworts have clearly recognisable leaves on clearly recognisable
stems but they totally lack a root system. All hornworts and some liverworts lack even a leaf–stem
differentiation but instead grow as ribbonlike thalli.” Dan Norris of University of California has the
largest collection of bryophytes in the world, which includes over 1,06,000 specimens.
1.1.1 Are all Mosses Bryophytes?

No, all “mosses” are not bryophytes. In bryophyta, all mosses belong to the class Musci or Bryopsida,
which includes more than 600 genera and more than 15,000 species. They have a leafy (not thalloid)
gametophyte, multicellular rhizoids, and a majority of them have a capsule with both a columella and a
lid or operculum. Chondrus crispus, an alga, is commonly called Irish moss while Cladonia rangifera
(a lichen) is commonly called Reindeer moss and Lycopodium (a pteridophyte) is commonly called
club-moss, and Spanish moss (Tillandsia usneoides) is a flowering plant.
1.1.2 What are Liverworts?

The word liverwort is a misnomer, and in bryophytes it is used for members of a class (Hepaticae or
Hepaticopsida). Liverworts are both thalloid as well as leafy. Some thalloid members of Hepaticopsida
resemble the liver, and hence the name “liverwort” is given. There is, however, no evidence that these
liverworts are of medicinal value. In thalloid liverworts, the gametophyte is a flat, more or less
undifferentiated thallus, whereas in leafy liverworts, the gametophyte has a simple stem, growing from
the apex and bearing small leaves in the rows along it. About 80% of the liverwort species are leafy,
while only about 20% of liverwort species are thalloid.
Introduc on � 3
HOW DO RECENT SYSTEMATISTS TREAT THE TERMS

“BRYOPHYTA”, “BRYOPHYTES” AND “BRYOBIOTINA”? 1.2
On the basis of recent genetic information, available molecular data between 2001 and 2010, and the
current concepts of phylogeny and classification, systematists have started treating these terms quite
differently from their existing explanations.
1. The hornworts, sharing their small size and independent, dominant gametophyte and dependent
sporophyte with the mosses and liverworts, have been considered by most systematists and
recent bryologists now to be in a separate phylum, the ANTHOCEROTOPHYTA.
2. A majority of the recent bryologists now also agree that the liverworts should occupy an
independent phylum, the MARCHANTIOPHYTA (also known variously as Hepatophyta,
Hepaticophyta, Hepaticopsida and Hepaticae).
3. Only mosses (included earlier under the class Musci) are now treated as the only members of
the phylum BRYOPHYTA.
4. All mosses, liverworts and hornworts together are still considered by bryologists under the
English name BRYOPHYTES, a term having no taxonomic status, and some bryologists have
suggested the rank of a subkingdom under the name BRYOBIOTINA.
All bryophytes are, therefore, now treated by recent bryologists (Crum, 2001; Norris, 2003; Shaw
and Renzaglia, 2004; Zander, 2006; and Troitsky et al., 2007) under the subkingdom BRYOBIOTINA,
which are classified into three phyla, viz. Marchantiophyta ( = liverworts), Bryophyta ( = mosses)
and Anthocerotophyta ( = hornworts). The characters, which all bryophytes share with tracheophytes,
are:
1. Development of an embryo within a multicellular reproductive organ
2. Presence of a covering of sporopollenin on their spores
3. Presence of flavonoids.
The characters in which all bryophytes differ from tracheophytes are the following:
1. Bryophytes have a dominant gametophyte supporting a parasitic sporophyte.
2. They lack meristematic tissue, lignin, tracheids and sieve cells.
Only due to the characters, like lack of lignin, and lack of tracheids and vessels, bryophytes have a
very small stature and are also termed nontracheophytes.
WHY ARE BRYOPHYTES ALWAYS SMALL SIZED

AND LACK TRUE LEAVES? 1.3
Bryophytes are small-sized plants, and a majority of them attain a length or height of only up to a few
centimetres. According to Hebant (1977), “one contribution to their small size is their lack of lignin,
limiting their size to that which their unlignified tissues can support.” However, there are conflicting
reports about the presence or absence of lignin in bryophytes. Downey and Basile (1989) reported lignin
in the sporophytes of Pellia epiphylla, and Crum (2001) reported lignin-like compounds in peristomes
of some bryophytes. However, the presence of lignin in gametophytes of bryophytes has still not been
conclusively evidenced. A lignin-like substance in the cell walls of Rhacocarpus purpurascens, a
moss, has been reported by Edelmann et al. (1998). Some phenolic compounds, similar to lignin, have
certainly been reported in many bryophytes.
4 � Bryophyta
The characteristic of “being without lignin” imposes some other limits on the plants. Due to absence
of lignin, bryophytic plants have no tracheids or vessels, hence they lack the type of the conducting
system, which is present in tracheophytes or vascular plants. This implies that the bryophytes lack true
vessels, and are also, therefore, called nontracheophytes.
Instead of tracheids or vessels, many bryophytes possess hydroids that confer much the same
function as xylem. Many others possess leptoids, the moss version of phloem. Many moss stems
possess a central strand, with or without hydroids, but with elongate cells. It functions in conduction.
Because of the greater density of smaller cells in the conducting strand, it also provides support to the
plant.
WHY DO BRYOPHYTES POSSESS RHIZOIDS

INSTEAD OF ROOTS? 1.4
Bryophytes lack tracheids and vessels and also, therefore, a sophisticated tracheid conducting system.
This limits or slows the movement of water within the plant, and, therefore, the roots are absent
in bryophytes. This lack of roots is substituted in most bryophytes by the nonvascular thread-like
structures, called rhizoids.
Rhizoids in bryophytes help in obtaining nutrients from a larger soil volume. They also help in
slowing the process of desiccation. Due to this characteristic, many bryophytes are desiccation tolerant.
NUMBER OF SPECIES AND GENE BANK DATABASE 1.5

As mentioned in some more detail in Chapter 2, in modern classifications, bryophytes are divided into
three groups having the “rank or divisions: Bryophyta, Marchantiophyta and Anthocerotophyta, or even
superdivisions” (Kenrick and Crane, 1997; Troitsky et al. 2007). And according to estimates of Ignatov
(2007), modern bryophytes—hornworts ( = Anthocerotophyta), liverworts ( = Marchantiophyta) and
mosses ( = Bryophyta)—“include about 100, 5,000 and 10,000 species, respectively”.
According to Troitsky et al. (2007), “As of September 2007, there are 337 entries for hornworts,
5517 for liverworts, and 17412 for mosses in the GeneBank database.” The current state of the
problem with the number of bryophyte species and their GeneBank database is reflected in Molecular
Systematics of Bryophytes by Goffinet et al. (2004) and in an article of Renzaglia et al. (2007) in the
journal Bryologist.
AGE OF BRYOPHYTES 1.6

As detailed in a review entitled “Contribution of Genosystematics to Current Concepts of Phylogeny
and Classification of Bryophytes”, Troitsky et al. (2007) stated that “The time of origin of bryophytes
and their phyla is not well determined from paleontological data. The cause is a bad preservation of
these plants in sediments and rarity of findings of sporophytes, on which the modern systematics is
greatly based. Most of such well-identified forms are <60 million years old. However, the age of fossil
spores, which can be classified as bryophytes, is 440–450 million years, which is in good agreement
Introduc on � 5
with the most reliable molecular-genetic chronology of the origin of land plants (425–490 million years
ago).”
OCCURRENCE AND DISTRIBUTION 1.7

Represented by approximately 960 genera and over 15,000 (Gradstein et al., 2001) to 25,000 species
(Crum, 2001), bryophytes are of widespread occurrence in almost all parts of the world. They grow
in almost all habitats but are mainly amphibious in nature. A majority of the bryophytes are moisture-
loving terrestrial plants growing in shaded grounds, on moist rocks, trunks of trees and other similar
moist places. They usually grow in tufts and cushions, and are responsible for providing the green colour
to the grounds, forests and mountains, especially during the rainy season. Bryophytes grow luxuriantly
in humid climates of both tropical and temperate regions. Liverworts grow commonly in humid tropics
but are rare in arctic environments. Mosses, however, survive in arctic and alpine regions. Only a few
bryophytes are aquatic, e.g. Riccia fluitans, Ricciocarpos natans, Riella sp. and Fontinalis antipyretica.
A few members (e.g. Leucobryum glaucum, Calypogeia fissa and Sphagnum sp.) grow in bogs, i.e.
wet and soft grounds, while some are found in xeric conditions, e.g. Torula muralis and Polytrichum
juniperinum. Buxbaumia and Cryptothallus mirabilis are saprophytic bryophytes.
Though bryophytes are not found in the sea, some mosses (e.g. Grimmia maritima and Eurhynchium
praeolongum) grow in crevices of rocks regularly bathed by sea water. Bryophytes usually occur on
the mountains at an altitude of 4000-8000 feet but a few mosses (e.g. Aongstroemia julacea) have also
been reported from an altitude of even 20,000 feet.
GAMETOPHYTE PLANT BODY 1.8

The plant body is gametophytic, and the gametophyte is independent and autotrophic in bryophytes. It
is either thalloid (i.e. thallus-like, and not differentiated into roots, stem and leaves; (Fig 1.1a) or foliose
(i.e. containing a definite rootless “leafy” shoot; Fig 1.1b). The gametophytic phase is, therefore, the
dominant and longer-lived phase of the life-cycle as compared with the sporophytic phase. According
to Renzagalia et al. (2000), the “bryophyte gametophytes are amongst the most elaborate of any phylum
of plants”.
The plant body of most bryophytes is small-sized, reaching only up to a few centimetres in length.
However, some species of the Australasian genus Dawsonia reach up to a height of 40–70 cm. Fontinalis
antipyretica, a moss, attains a length of 50–70 cm, while Martin (1951) recorded a remarkable example
of a stem of Polytrichum commune attaining a length of over 180 cm. Crum (2001) mentioned that
some mosses (e.g. Ephemeropsis and Viridivellus pulchellum) are only a few millimetres tall and have
only a few leaves. The liverwort thallus of Monocarpus is only 0.5–2 mm in diameter, and some
Fontinalis species can be 2 m in length.
True roots, stems and leaves, as found in vascular plants, are completely absent in the plant body
of bryophytes. The “stem” and “leaflike” structures of foliose bryophytes have been termed “axis”
and “phylloid” by Koch (1956). Vascular tissues (xylem and phloem) are completely absent in all
bryophytes and, therefore, Tippo (1942) suggested the name Atracheata to them. Cuticle and stomata
are also absent.
6 � Bryophyta
Fig. 1.1 (a) Thalloid plant body of Riccia (b) Foliose plant body of Funaria
The plant bodies of thalloid members (e.g. Riccia, Marchantia, Anthoceros) grow prostrate on the
ground and remain attached to the substratum with the help of several fine, delicate, hairlike, unicellular
organs called rhizoids. Along with the rhizoids, several scales may also be present in several genera
(Riccia, Marchantia) on the ventral surface of the thalloid plant body. Sex organs, i.e. antheridia and
archegonia, are present on the dorsal surface of thalloid genera.
The plant body of foliose members (i.e. leafy Hepaticopsida and mosses) usually grows erect and
remains differentiated into rhizoids, axis (comparable to stem) and leaflike structures called phylloids
or phyllids. The rhizoids develop from the basal older parts of the axis and are usually well-branched
and multicellular.
Developing on the gametophyte, and dependent on it for nutrition and physical support, is the
sporophyte. It is usually differentiated into foot, seta and capsule (Fig. 1.1b). The foot is usually
embedded within the gametophyte and functions for anchorage and absorption. Seta is a long stalk-
like structure and helps in projecting the capsule, which is a spore-producing body of the sporophyte.
The sporophyte has no connection with the soil and is completely dependent on the gametophyte for its
water and mineral nutrition supply.
VEGETATIVE REPRODUCTION 1.9

Vegetative reproduction is the most prevalent and efficient method of reproduction in most bryophytes.
It takes place by a variety of ways, described briefly as under:
Introduc on � 7
1. Fragmentation, due to the progressive death and decay of older parts, as in majority of
Hepaticopsida and Anthocerotopsida.
2. Adventitious branches, often developing from the underside of the midrib, as in Riccia
fluitans, Marchantia, Corsinia, Reboulia, Dumortiera and Anthoceros.
3. Innovations, which are the axillary branches, growing vigorously in some plants, such as
Sphagnum and several acrogynous Jungermanniales.
4. Phylloid cladia, which are small detachable branches, originating from the individual cells of
the ‘leaf’ or phylloid, as in Bryopteris and Frullania.
5. Axis cladia, which originate in the axis in some foliose genera and occupy the same position
as the sexual branches, as in Leptolejeunea.
6. Tubers, which develop in the form of swollen tips of the underground branches during drought
and other unfavourable conditions, are formed in several species of Riccia, Anthoceros,
Fossombronia, Asterella, Conocephalum, etc. Food reserves, such as starch grains and oil
globules, remain packed in tubers.
7. Gemmae, the propogative organs of definite form, are formed in several genera of
Hepaticopsida, Anthoceropsida and Bryopsida. Gemmae are large, stalked and multicellular
in Marchantia and Lunularia; small, multicellular and discoid in Radula; bicelled, endogenous
bodies in Riccardia multifida; discoid and platelike in Metzgeria; filamentous, microscopic
and multicellular in Torula papillosa and, globular and multicellular in Bryum rubens.
8. Primary protonema, which develop by the germination of spores, as in Funaria.
9. Secondary protonema, which develop from the methods other than the germinating spores,
as in Sphagnum and Funaria.
10. Apospory which is the production of diploid gametophyte from the vegetative cells of
sporophyte, i.e. without the production of spores, as in Anthoceros and several mosses.
Vegetative reproduction in bryophytes has been discussed in detail in Chapter 18.
SEXUAL REPRODUCTION 1.10

Without exception, the sexual reproduction in all bryophytes is of highly oogamous type, i.e. takes
place by motile male gametes (antherozoids) and a large nonmotile female gamete (egg). The gametes
are produced within the multicellular sex organs, which are provided with an outer sterile layer of
jacket. Such an outer layer of sterile jacket is absent in the sex organs of algae. The male sex organ is
called antheridium while the female sex organ is called archegonium.
Antheridium (Fig. 1.2) is a multicellular, ellipsoidal or club-shaped, short-stalked body consisting
of a mass of androcytes or antherozoid mother cells surrounded by a single layer of protective sterile
jacket cells. Each androcyte produces a single biflagellate motile male gamete called antherozoid.
Usually, the antherozoids are spirally coiled bodies, each possessing two whiplash-type flagella. In
most bryophytes, the antheridium, on dehiscence, discharges the androcytes in a mass.
The archegonium (Fig. 1.3) is also a multicellular but flask-shaped body with a basal swollen
portion called venter and an upper elongated portion called neck. Each archegonium consists of an
axial row of cells surrounded by a sterile jacket. The axial row of archegonium contains usually 4–6
or more neck canal cells, a venter canal cell and a single large basal cell called egg or oosphere.
Nourishment and protection to the egg and the developing embryo are provided by the archegonium.
8 � Bryophyta
Jacket
Androgonial
cells
Stalk
Fig. 1.2 A mature antheridium of Riccia Fig. 1.3 A mature archegonium of Riccia
Fertilization takes place only in the presence of water. At the time of fertilization, the axial row
of neck canal cells and venter canal cell of the archegonium disintegrate and disorganise (Fig. 1.4),
forming a mucilaginous liquid. Only the egg remains inside the cavity of the venter. The liberated,
flagellated, antherozoids swim in a thin film of water and reach up to the neck of the archegonium. The
mouth or the tip of the archegonium also opens at this stage. Several antherozoids pass through the neck
canal to the venter, where a single antherozoid fertilizes the egg, and a diploid zygote is resulted.
Gametes (antherozoids and egg) are the last structures of the gametophytic generation, while the
zygote is the first cell of the sporophytic generation.
Fig. 1.4 An archegonium of Riccia just before fer liza on

Introduc on � 9
SPOROPHYTE 1.11
The diploid zygote is the starting point of the sporophytic generation. The zygote, without any resting
period, begins to grow at once, divides and redivides, and forms a multicellular embryo. The embryo
develops into a sporophyte or sporogonium. The sporophyte in bryophytes is never an independent
body. It is always dependent on the parent gametophytic plant. The wall of the venter enlarges with the
developing embryo to form a protective covering called calyptra. The multicellular embryo obtains
its nourishment directly from the gametophytic thallus, to which it is attached. The young embryo is
an undifferentiated mass of cells. But it soon gets segmented and differentiated into well-developed
sporophyte.
The sporophyte is usually differentiated into three parts, viz. foot, seta and capsule (e.g. Marchantia,
Porella, Fig. 1.5). In certain cases, the seta is absent (e.g. Corsinia), and in certain others, both seta and
foot are absent (e.g. Riccia). The foot is basal and usually a bulbous structure, embedded in the tissue of
the gametophyte. Its function is absorption. The seta is present in between the foot and the capsule. Its
main function is elongation. It also conducts the food absorbed by the foot to the capsule. The capsule
is the terminal part of the sporophyte and its function is spore production. Inside the capsule are present
some sterile elater mother cells and fertile spore mother cells. Elater mother cells change into long,
slender elaters, which are hygroscopic in nature. The spore mother cells are diploid and represent
the last stage of sporophytic generation. They divide reductionally to form spore tetrads, which soon
separate into haploid spores or meiospores. Each spore develops into a young haploid gametophyte.
Calyptra
Capsule
Elaters
Spore
Seta
Foot
Thallus cells
Fig. 1.5 A mature sporophyte of Porella

10 � Bryophyta
YOUNG GAMETOPHYTE 1.12

The spore is the initial stage of the gametophytic plant body. The spores are usually morphologically
similar in shape, size and structure, i.e. they are homosporous in bryophytes. Each spore is unicellular
and haploid, and germinates into a young gametophytic plant (e.g. Riccia, Marchantia). In Funaria and
several other mosses, the spore first germinates into a filamentous protonema, from which the buds are
produced to give rise to young gametophytic plants.
LIFE CYCLE ALTERNATION OF GENERATIONS 1.13

Bryophytes are peculiar in that their life-cycle is completed by a regular alternation between two
generations: a gamete-producing generation (gametophyte) and a spore-producing generation
(sporophyte). Because both of them are quite distinct from each other and also are quite dissimilar in
their sizes, they show heteromorphic alternation of generations.
In the life-cycle of these plants, there exist two distinct phases. One is the haploid (X) or gametophytic
phase, with one complete set of chromosomes in the cells. This phase is represented by thalloid (e.g.
liverworts) or foliose (e.g. mosses) gametophytic plant body, which is the dominant and independent
phase of the life-cycle. The gametophytic plant body is haploid and bears the sex organs, i.e. antheridia
and archegonia. Haploid gametes are produced in these sex organs, i.e. antherozoids in the antheridium
and egg in the archegonium. (The green individual plant is called ‘gametophyte’ since it bears the
gametes). The haploid or gametophytic generation is also called sexual generation, as it bears the sex
organs. Two different gametes fuse under the process of fertilization and form a diploid zygote. The
zygote is the starting point of the next phase of the life-cycle, i.e. diploid (2X) or sporophytic phase.
On germination, the zygote does not produce the gametophytic plant. It divides and redivides to form
a multicellular embryo, which, by further segmentation, is differentiated into the second diploid adult
of the life-cycle called sporophyte. In the capsular region of the sporophyte are present the diploid
spore mother cells, which undergo meiosis and form haploid spores called meiospores. The zygote,
embryo, sporogonium and spore mother cells together constitute the sporophytic generation. This is
called ‘sporophytic generation’ because it is concerned with the production of spores. Since two sets
of chromosomes exist in the cells of all stages of this phase, it is also called diploid phase (2X). It is
also called non-sexual generation because it ends in the production of asexual spores. The sporophytic
generation is dependent, usually completely or sometimes partially, on the gametophytic generation
for its nutrition. The haploid spores, produced at the end of this diploid phase, are the starting point
of the haploid gametophytic plant body. Each spore germinates and produces a gameteophytic plant
body. Each spore germinates and produces a gametophytic plant, which again bears the sex organs
(antheridia and archegonia). And in this way, the life-cycle goes on. Since the sex organs, i.e. antheridia
and archegonia, are not exposed, these plants are described as cryptogams (Greek—kruptos, hidden;
gamos, wedded). Because the two generations (gametophytic and sporophytic) appear alternately in the
life-cycle, these plants show alternation of generations. Since the two generations differ completely
in their morphology (i.e., gametophyte is either thalloid or foliose, and the sporophyte usually consists
of foot, seta and capsule), it is called heteromorphic alternation of generations.
Introduc on � 11
Fig 1.6 Diagramma c representa on of the life cycle in bryophytes
A NOTE ON REGENERATION IN BRYOPHYTES 1.14

Bryophytes have exceptional capacity of regeneration. In the dry conditions, the gametophytic plant
body of these plants becomes brittle and its isolated fragments readily regenerate to form entire plants.
Regeneration is also shown by the cells of the sporophyte. These cells regenerate to develop into a
protonema, and soon the gametophytes appear on these protonemal bodies. Such a regeneration of
gametophytes from the sporophytic cells, without the formation of spores, is called apospory. On
the contrary, in some bryophytes there is seen the regeneration of a sporophyte from a gametophyte,
without the formation of gametes, and this phenomenon is called apogamy. Gemmae (cluster of cells
on the gametophyte, which, on separation from the parent plant, give rise to a new gametophytic plant)
are also the effective means of regeneration in several bryophytes, e.g. Marchantia. In bryophytes,
however, the phenomena of apospory and apogamy are not of common occurrence.
SALIENT FEATURES OF BRYOPHYTES: AT A GLANCE 1.15

1. Bryophytes live in wet, shady habitats and include liverworts, hornworts and mosses.
2. Their vegetative plant body is completely adapted to the land habit. For their sexual
reproduction, however, they rely upon water because their swimming habit is retained by their
male gametes or antherozoids.
12 � Bryophyta
3. Their plant body is gametophytic, which is independent.

4. The plant body lacks true roots, stem and leaves.
5. It is either thalloid or foliose.
6. Most bryophytes have very little or no vascular system. The characteristic vascular tissue
(xylem and phloem) of higher plants is absent.
7. Thalloid bryophytes (e.g. Riccia, Marchantia) grow prostrate on the ground and remain
attached to the substratum by several delicate, unbranched, unicellular, hairlike structures
called rhizoids.
8. Foliose bryophytes (e.g. Funaria) have an erect plant body possessing a central axis bearing
many leaflike expansions. They remain attached to the substratum by many branched,
multicellular rhizoids.
9. Gametophyte is concerned with the sexual reproduction, which is invariably highly
oogamous.
10. Differing from algae, the sex organs in bryophytes are jacketed and multicellular.
11. The antheridium is club-shaped and multicellular, and bears many biflagellate antherozoids,
of which both the flagella are of whiplash type.
12. The archegonium is the female sex organ. It is a flask-shaped body, containing a globular
venter and long neck. The female gamete is in the form of an egg.
13. Fertilization takes place only in the presence of water.
14. The fertilized egg or zygote is retained within the venter of the archegonium and divides only
there. It never becomes independent, i.e. it always remains within the parent gametophyte.
15. Repeated divisions of the zygote results into the formation of a multicellular embryo.
16. The first division of zygote is transverse, and the apex of the embryo develops from the outer
cell. The embryogeny is thus exoscopic, a characteristic feature of bryophytes.
17. The venter of the archegonium develops into a protective, multicellular structure called calyptra.
18. Embryo, in the later stages, differentiates into a spore-producing structure called sporophyte
or sporogonium.
19. The sporophyte in bryophytes is never differentiated into structures like roots, stem and leaves.
It is always dependent, partially or completely on the gametophyte, and usually contains
structures like foot, seta and capsule.
20. Diploid spore mother cells of the sporogonium divide reductionally to form haploid spores or
meiospores.
21. Meiospores of one species are morphologically similar. Bryophytes are, therefore,
homosporous.
22. Each spore germinates into a haploid gametophytic plant.
23. All bryophytes show heteromorphic alternation of generations.
AFFINITIES OF BRYOPHYTES 1.16

Bryophyta are a group of amphibian plants occupying a position in between algae (being aquatic) and
pteridophytes (being terrestrial). They, thus, have some affinities with both algae and pteridophytes,
discussed briefly as follows:
Introduc on � 13
1.16.1 Affini es of Bryophytes with Algae

1. Resemblances of Bryophytes with Algae
(a) Both bryophytes and algae have thallus-like bodies.
(b) Both lack vascular tissue, i.e. xylem and phloem.
(c) Roots are absent in both.
(d) Visible plant body in all bryophytes and most algae is gametophytic.
(e) Members of both groups possess chlorophyll and are thus autotrophic.
(f) Chlorophyll a and b, xanthophyll and carotene are present in both bryophytes and in green
algae (Chlorophyta).
(g) Antherozoids of both groups have a swimming habit.
(h) Antherozoids of both groups have whiplash flagella.
(i) Filamentous thalli of many green algae show striking similarities with the early developmental
stages of gametophytes of many bryophytes.
(j) In both bryophytes and green algae, the carbohydrate food is a true starch, containing a mixture
of two kinds of glucose macromolecules, amylose and amylopectin.
(k) In both bryophytes and green algae, the structure and composition of cell walls show many
resemblances. They possess cellulose cell walls.
(l) Presence of pyrenoids in green algae and Anthocerotales (e.g. Anthoceros) is also a striking
similarity between them.
The resemblances mentioned above support the theory of the evolution of bryophytes from algae,
especially from the green algae (Chlorophyceae).
2. Differences between Bryophytes and Algae

Some of the striking differences between algae and bryophytes are listed in Table 1.1.
1.16.2 Affini es of Bryophytes with Pteridophytes

1. Resemblances of Bryophytes with Pteridophytes
(a) Terrestrial habit is usually shown by both groups.
(b) Asexual spores (mitospores) are absent in the life-cycle of members of both groups.
(c) Oogamous sexual reproduction is shown by members of both groups.
(d) Plants are archegoniate in both groups, and structure of archegonium in both groups is almost
similar.
(e) A layer of sterile jacket cells surrounds the antheridium in both.
(f) Male gametes are flagellated in members of both groups.
(g) Water is essential for fertilization in both.
(h) All members of both the groups show development of zygote into an embryo.
(i) In both groups, the embryo develops within the archegonium protected by multicellular
maternal tissue.
(j) A well-marked heteromorphic alternation of generations is shown by members of both
groups.
14 � Bryophyta
Table 1.1 Differences between algae and bryophytes
S. NO. ALGAE BRYOPHYTES

1 Most algae are aquatic. Chiefly terrestrial, occurring in moist shady
places.
2. Plant body is either unicellular or multicellular, Plant body is multicellular and thalloid or foliose
filamentous or pseudoparenchymatous. (differentiated into rhizoids, axis and leaflike
structures).
3. Filaments are unbranched or branched, showing Thalloid plant body is usually dichotomously
irregular branching. branched.
4. Rhizoids are usually absent except in some het- Rhizoids are present in all bryophytes.
erotrichous forms.
5. Almost all cells of the plant body are capable of Divisions are restricted only to certain apical
divisions. cells.
6. Asexual reproduction by formation of mito- Mitospores, like zoospores, are completely
spores, e.g. zoospores. absent.
7. Sexual reproduction is isogamous, anisogamous Sexual reproduction is always oogamous.
or oogamous.
8. Sex organs do not possess a covering of sterile Sex organs are always covered by a single sterile
cells. layer of jacket.
9. The female sex organ is usually a unicellular Female sex organ is always a multicellular
oogonium. archegonium.
10. The diploid zygote is usually liberated from the The zygote is never liberated. It starts dividing
female sex organ and only then it starts dividing. into a sporogonium inside the gametophytic
plant body only.
11. Prior to germination, the zygote requires a resting The zygote does not require a resting period. It
period. starts dividing immediately after its formation.
12. There is no embryo formation after gametic All bryophytes show development of zygote into
union. an embryo.
13. The diploid sporophytic phase is in the form of a The diploid sporophytic phase is never an
zygote which is an independent body. independent body. It is always dependent on the
gametophyte.
14. Alternation of generations, if present, is of ho- All bryophytes exhibit alternation of generations,
mologous type. and it is always of heteromorphic type.
15. Gametophytes and sporophytes are both indepen- Sporophyte in all bryophytes is always depen-
dent bodies. dent on gametophyte, either wholly or partially.
Introduc on � 15
2. Differences between Bryophytes and Pteridophytes

The major differences between bryophytes and pteridophytes are listed in Table 1.2.
Table 1.2 Differences between bryophytes and pteridophytes
S. NO. BRYOPHYTES PTERIDOPHYTES

1. Dominant phase in the life-cycle is gameto- Dominant phase in the life-cycle is sporophytic.
phytic.
2. Plant body is either thalloid or foliose. It is Plant body is always differentiated into roots, stem
never differentiated into roots, stem and leaves. and leaves.
3. Vascular tissue (xylem and phloem) is absent. Vascular tissue in the form of xylem and phloem is
always present.
4. Sporophyte is dependent on the gametophyte. Sporophyte is independent. It is never dependent on
the gametophyte.
5. All bryophytes are homosporous. Pteridophytes are both homosporous as well as
heterosporous.
ORIGIN OF BRYOPHYTES 1.17

Discussed in detail in Chapter 31.
TEST YOUR UNDERSTANDING
1. Who was the first to introduce the word “bryophyta”?

2. Bryophyta includes liverworts, hornworts and ___________.
3. The name “amphibians of the plant kingdom” is o en given to _________.
4. What are bryophytes? Explain in about 100 words.
5. Embryophyta includes _____, pteridophyta and spermatophyta.
6. Are all “mosses” bryophytes?
7. Differen ate between the terms “liverworts” and “hornworts”.
8. Discuss briefly how the modern systema sts treat the terms (a) Bryophyta, (b) Bryophytes,
and (3) Bryobio na?
9. Write at least 10 salient features of bryophytes.
10. Is vascular ssue completely absent in bryophytes?
11. Explain in brief various methods of the vegeta ve reproduc on in bryophytes.
12. Antheridia of bryophytes are club-shaped while their archegonia are ________ shaped.
13. Bryophytes are always small-sized and lack true leaves. Why?
14. How will you differen ate between the sex organs of algae and bryophytes?
15. Why do bryophytes possess rhizoids and not roots?
16. Write a brief note on the sporophyte in bryophytes.
17. Explain alterna on of genera ons in bryophytes.
18. Discuss affini es of bryophytes in detail.
2
Classification
CLASSIFICATION 2.1
Braun (1864), who introduced the term “Bryophyta”, included all algae, lichens and mosses under
bryophytes. The rank of the division to the Bryophyta was first given by Schimper (1879).
Eichler (1883) was the first to divide Bryophyta into two classes, viz. Hepaticae and Musci. Engler
(1892) also followed Eichler but divided the class Hepaticae into the following three orders:
1. Marchantiales
2. Jungermanniales
3. Anthocerotales
The abovementioned same system of classification of Bryophyta and Hepaticae has also been
followed by some eminent botanists including Fritsch (1929), Bower (1935), Evans (1939) and also in
Syllabus der Pflanzenfamilien by Engler, Melchior and Werdermann (1954).
Underwood (1894) and Gayet (1897) studied in greater detail the evolution of Hepaticae, and
suggested that there exist several fundamental differences between Anthocerotales and the remaining
members of Hepaticae. On the basis of such differences, several later workers (Smith, 1938, 1955;
Takhtajan, 1953; Wardlaw, 1955; and Schuster, 1958) divided Bryophyta into following three
classes:
1. Hepaticae
2. Anthocerotae
3. Musci
Howe (1899) was actually the first to provide a separate ‘class’ status to Anthocerotales and has
named the class ‘Anthocerotes’.
International Code of Botanical Nomenclature suggested in 1956 that the suffix —‘opsida’ should
be used for the classes, and all subclasses should end with the suffix- “idae”. And, such usage had
already been proposed by Rothmaler (1951) for the classes of Bryophyta. He suggested that the names
Hepaticopsida, Anthoceropsida and Bryopsida should be used instead of the older names, i.e.
Hepaticae, Anthocerotae and Musci for the classes of Bryophyta.
Classifica on � 17
Proskauer (1957) suggested that the class name Anthoceropsida should be changed to
Anthocerotopsida. Proskauer’s classification, followed by Parihar (1965), Holmes (1986) and most
other bryologists has been followed in this book. It divides Bryophyta into three classes:
1. Hepaticopsida
2. Anthocerotopsida
3. Bryopsida
An outline of the classification of the division Bryophyta, suggested by Sandra Holmes (1986) in her
book Outline of Plant Classification, is mentioned below:
Division BRYOPHYTA
Class 1 Bryopsida (Musci, mosses)
Subclass 1. Sphagnidae (peat or bog mosses)
Order—Sphagnales
Subclass 2. Andreaeidae (granite mosses)
Order—Andreaeales
Subclass 3. Bryidae (true mosses)
Cohort 1. Eubryidae
Orders—1. Archidiales 2. Dicranales 3. Fissidentales 4. Encalyptales 5. Pottiales 6. Grimmiales
7. Funariales 8.Eubryales (Bryales) 9. Orthotrichales 10. Isobryales 11. Hookeriales
12. Hypnobryales 13. Thuidiales 14. Schistostegales 15. Tetraphidales
Cohort 2. Buxbaumiidae
Orders—1. Buxbaumiales 2. Diphysciales
Cohort 3. Polytrichiidae
Orders—1. Polytrichales 2. Dawsoniales
Class 2. Hepa copsida (true liverworts)

Subclass 1. Jungermanniae
Orders—1. Takakiales 2. Calobryales 3. Jungermanniales 4. Metzgeriales
Subclass 2. Marchantiae
Orders—1. Sphaerocarpales 2. Monocleales 3. Marchantiales
Class 3. Anthocerotopsida (horned liverworts or hornworts)

Order—Anthocerotales.
MAJOR DIFFERENCES BETWEEN CLASSES OF BRYOPHYTA 2.2

Some basic differences between the three classes (Hepaticopsida, Anthoceropsida and Bryopsida) of
Bryophyta are given in Table 2.1:
18 � Bryophyta
Table 2.1 Basic differences between three classes of Bryophyta
S. NO. HEPATICOPSIDA ANTHOCEROPSIDA BRYOPSIDA

1. Spores are usually tetrahedral. Same as in Hepaticopsida. Spores are usually spherical.
2. Protonema is either absent or not Almost same as in Hepaticopsida. Protonema is well-developed. It is
sharply differentiated from game- sharply differentiated from game-
tophyte. tophytic plant body.
3. Gametophytic plant body is either Gametophytic plant body is thal- Plant body is gametophytic and
thalloid or foliose and strongly loid but in Dendroceros it becomes never thalloid. It is foliose.
dorsiventral. slightly foliose.
4. Leaf-like structures are present Leaflike structures are usually Leaflike structures always present.
only in foliose members. They are absent. They are always spirally arranged
never spirally arranged. If present, on the axis. A midrib is always
the leaves are always made of a present, except in some species of
single cell-thick plate of cells. A Andreaea and Sphagnum.
midrib is absent.
5. Rhizoids are thread-like un- Rhizoids are threadlike, un- Rhizoids are well-branched, mul-
branched, unicellular, and of two branched, unicellular and only ticellular. They usually contain
types, i.e. smooth-walled and smoothwalled. oblique septa.
tuberculate.
6. Scales are present in many mem- Scales absent. Scales absent.
bers, e.g. Riccia, Marchantia.
7. Antheridia are cylindrical or spher- Almost same as in Hepaticopsida. Antheridia are club-shaped, with
ical with no association of any hairs a massive stalk, and are always
or paraphysis-like structures. associated with many hairlike,
multicellular paraphyses.
8. Archegonia are usually sessile or Archegonia are completely sunk in Archegonia have a distinct stalk or
shortly-stalked, with a short neck; the gametophytic tissue; jacket of pedicel; neck is very long, twisted
jacket made up of 4–6 vertical rows neck is made up of 6 vertical rows and composed of 6 vertical rows
of cells; without any paraphyses. of cells; paraphyses absent. of cells; several paraphyses present
amongst the archegonia.
9. A four-celled lid or cover cells are Same as in Hepaticopsida. Cover cell functions as an apical
present at the top of each arche- cell for a long time and contributes
gonium. to the wall of the neck and the neck
canal cells.
10. Sporogonium is a simple structure, Sporogonium is not a simple Sporogonium is most complex
represented by only capsule (e.g. structure. A meristematic zone is amongst bryophytes, divisible into
Riccia) or divisible into foot, seta present in between a bulbous foot foot, seta and capsule, it shows a
and capsule (e.g. Marchantia). and spore-producing capsule. high degree of internal differentia-
tion of tissues.
11. Sporogonium growth is fixed and Sporogonium growth is continu- Sporogonium growth is indeter-
determinate. ous and indeterminate due to the minate; seta is meristematic in
presence of meristematic zone in nature.
between foot and capsule.
12. Columella is absent. Columella is usually present. Columella is mostly present.
13. Some genera bear elaterophore Elaterophore absent. Elaterophore absent.
(e.g. Pellia).
(Contd.)
Table 2.1 (Contd.)

S. NO. HEPATICOPSIDA ANTHOCEROPSIDA BRYOPSIDA
14. Archesporium or sporogenous Archesporium is usually Archesporium develops mostly
tissue is usually endothecial in amphithecial in orgin, with a few from the outermost layer of endoth-
origin. exceptions as in some species of ecium, except in a few cases as in
Notothylas. some species of Sphagnum.
15 Elaters (sterile filaments or cells Pseudoelaters are present instead Elaters or pseudoelaters are ab-
mixed with spores) are present in of elaters. sent.
the capsules of many genera.
16. Capsule dehisces either irregularly Capsule dehiscence is simple and Capsule dehiscence is usually
or by transverse or longitudinal takes place by 2 to 4 longitudinal brought about by operculum
slits. slits. and annulus, and it is the peris-
tome which controls dispersal of
spores.
LATEST POSITION OF CLASSIFICATION OF BRYOPHYTES 2.3

On the basis of the current state of morphology, available data of molecular studies, genosystematics,
phylogeny, diversification and classification of bryophytes, almost all modern bryologists (including
Clarke and Duckett, 1979; Smith, 1982; Schuster, 1984; Rykovskii, 1987; Newton et al., 2000;
Shaw and Goffinet, 2000; Crum, 2001; Norris, 2003; Shaw and Renzaglia, 2004; Zander, 2006; and
Troitsky et al., 2007)now agree that all bryophytes should be treated under an independent subkingdom
(BRYOBIOTINA), divisible further into the following three phyla:
1. Marchantiophyta ( = Liverworts or Hepatophyta, Hepaticophyta, Hepaticae, or
Hepaticopsida)
2. Bryophyta ( = Mosses or Musci or Bryopsida)
3. Anthocerotophyta ( = Hornworts or Anthoceropsida or Anthocerotae)
The phylum Bryophyta now, therefore, includes only “mosses” in the modern classifications.
Crum (2001), on the basis of morphological evidence, coupled with some biochemical evidence,
suggested the creation of a fourth phylum Sphagnophyta (including Sphagnum) under the subkingdom
Bryobiotina. However, several other workers (Rykovskii, 1987; Newton et al., 2000) supported the
concept of monophyletic origin for the bryophytes, including Sphagnum, when they studied data
from morphological, developmental, anatomical, ultrastructural and nucleotide sequence characters
together.
2.3.1 Marchan ophyta ( = Liverworts)

On the basis of available molecular data and studies of genosystematics, a completely new system of
classification of liverworts has been proposed by HeNygren et al. (2006). As detailed in the journal
Cladistics, an outline of this system is mentioned here as under:
20 � Bryophyta
Phylum Marchan ophyta
Class 1 Treubiopsida
Subclass: Treubiidae
Order—Treubiales
Subclass: Haplomitriidae
Order—Haplomitriales
Class 2 Marchan opsida

Subclass: Blastidae
Order—Blastiales
Subclass: Marchantiidae
Orders—Sphaerocarpales, Marchantiales
Class 3 Jungermanniopsida
Subclass: Pelliidae
Orders—Pelliales, Fossombroniales
Subclass: Metzgeriidae
Order—Metzgeriales
Subclass: Jungermanniidae
Orders—Pleuroziales, Porellales, Jungermanniales
2.3.2 Bryophyta (= Mosses)

The latest system, which has assimilated the genosystematics data, has been proposed by Goffinet and
Buck (2004) in Molecular Systematics of Bryophytes. It divides mosses into eight classes as under:
1. Takakiopsida
2. Sphagnopsida
3. Andreaeopsida
4. Andreaeobryopsida
5. Oedipodiopsida
6. Polytrichopsida
7. Tetraphidopsida
8. Bryopsida (largest class of mosses)
Troitsky et al. (2007) studied the phylogenetic relationships between basal groups of mosses on the
basis of molecular data and shown them as under:
2.3.3 Anthocerotophyta ( = Hornworts)

On the basis of methods of molecular phylogenetics, Duff et al. (2004, 2007) divided Anthocerotophyta
into the following two subclasses:
1. Anthocerotidae
2. Notothylatidae
1. All the bryophytes are tradi onally divided into three classes. Name them.
2. Who was the first to use the suffix - “opsida” for the classes of Bryophyta?
3. Proskauer’s classifica on of Bryophyta has been followed by most of the later workers,
including Parihar (1965) and Holmes (1986). Give an outline classifica on of Bryophyta used
by Parihar.
4. Write any seven major differences between Hepa copsida, Anthoceropsida and Bryopsida.
5. How do modern bryologists classify bryophytes, now in the 21st century?
3
Hepaticopsida:
Takakiales and
Calobryales
WHAT ARE HEPATICOPSIDA = LIVERWORTS ? 3.1
Formerly known as Hepaticae or “liverworts”, Rothmaler (1951) suggested the name “Hepaticopsida”
to the class taxon of bryophytes. The names (“Hepaticopsida” for Hepaticae, “Anthoceropsida”
for Anthocerotae and “Bryopsida” for Musci) have also been recognised by ICBN. Recently, some
taxonomists have started calling Hepaticopsida as “Marchantiopsida”.
In The Penguin Dictionary of Botany, Hepaticopsida or Hepaticae (Marchantiopsida) is “a class
of Bryophyta containing the thallose and leafy liverworts, which number about 10,050 species in
about 295 genera”. They “differ from the Musci (mosses) in showing marked dorsiventrality in the
gametophyte. The antheridia and archegonia may be borne on the surface of the thallus or on fleshy
stalks (gametangiophore). The capsule of the sporophyte, which contains sterile elaters as well as
fertile spores, matures before the seta lengthens, while in mosses the reverse occurs. The capsule does
not contain a central pillar of the sterile cells (columella) as is found in Musci and Anthocerotae.”
In the Chambers Biology Dictionary, Hepaticopsida is described as a class of Bryophyta, in which
“the gametophyte is thalloid or leafy with unicellular rhizoids and the capsule (sporophyte) without a
columella”.
WHY ARE THEY CALLED LIVERWORTS OR HEPATICOPSIDA? 3.2

In earlier times, it was believed that plants resembling the shape of the liver were good for liver ailments.
Since many bryophytes were thalloid or liverlike in their external morphology, the name Hepaticae or
Hepaticopsida was given to them, because the Latin word for liver was hepatica. Only because of this,
the common name “liverworts” was given to these plants. It is, however, not correct that all liverworts
are good for liver ailments.
Hepaticopsida: Takakiales and Calobryales � 23
GENERAL CHARACTERISTICS 3.3

1. The vegetative plant body is gametophytic and the gametophyte is dorsiventral and either
thalloid (i.e. thallus like) or foliose (i.e. bearing a leafy axis).
2. In foliose members, the leaves are arranged on the axis either in two or three rows.
3. A well-marked midrib is almost absent in the leaves.
4. Photosynthetic cells of the gametophyte contain numerous chloroplasts, which lack
pyrenoids.
5. One to many refractive oil bodies are usually present in all the chlorophyll-containing cells.
6. The sex organs are in the form of well-developed antheridia and archegonia.
7. Each sex organ develops from a single initial cell.
8. Fertilization results in the formation of a sporogonium.
9. The sporogonium is usually devoid of chlorophyll-containing cells.
10. Stomata are also absent in sporogonium.
11. The sporogonium is either simple (e.g. Riccia), or differentiated into foot, seta and capsule
(e.g., Marchantia). In some members, it is made of only foot and capsule with no seta.
12. The archesporium (the tissue that gives rise to the spore mother cells) is endothecial in
origin.
13. The sporogenous tissue of the capsule gives rise to fertile spore mother cells and sterile elaters
(e.g. Marchantia). In a few genera, the elaters are absent (e.g. Riccia).
14. In most of the genera, the elaters are unicellular structures with spiral thickenings in their
walls.
15. Columella is absent in the capsule of the sporogonium.
16. The capsule generally remains covered by a well-developed jacket. On being dry, the jacket
ruptures releasing the spores.
CLASSIFICATION OF HEPATICOPSIDA 3.4

Tracing history of the classification of Hepaticopsida, Engler (1892) included only two orders, namely
Marchantiales and Jungermanniales in this class.
Due to the presence of a special envelope around sex organs in members of the family
Sphaerocarpaceae of order Jungermanniales, this family has been elevated to the rank of an order
Sphaerocarpales by Cavers (1910). Class Hepaticopsida has thus been divided into three orders—
Marchantiales, Jungermanniales and Sphaerocarpales (Cavers,1910). Division of this class into the
same three orders has also been favoured by bryologists like Evans (1939), and Engler, Melchior and
Werdermann (1954).
Campbell (1936) separated the family Calobryaceae from the order Jungermanniales and proposed
to elevate it as a separate order Calobryales on the basis of peculiar characters like
1. Somewhat erect stems, bearing three rows of similar leaves, and
2. One-celled-thick jacket of the capsule. Four orders are thus recognised in Hepaticopsida:
(i) Marchantiales, (ii) Jungermanniales, (iii) Sphaerocarpales, and (iv) Calobryales.
Instead of Hepaticopsida, Schuster (1953, 1958) preferred to call this class as Hepaticae and divided
it into two subclasses: (i) Jungermanniae, including three orders, namely Calobryales, Jungermanniales
24 � Bryophyta
and Metzgeriales, and (ii) Marchantiae, including two orders, namely Sphaerocarpales and Marchantiales.
Schuster, on the basis of his detailed study of a large number of Hepaticae from USA, New Zealand
and Tasmania, thus divided this class into five orders, viz. (i) Calobryales, (ii) Jungermanniales,
(iii) Metzgeriales, (iv) Sphaerocarpales, and (v) Marchantiales.
The unigeneric family Monocleaceae shows characters of both Marchantiales and Jungermanniales
and has been placed by some bryologists amongst Marchantiales, and by others amongst Jungermanniales.
Monoclea, however, shows Calobryum-type of archegonial development, thus showing its closeness
with Calobryales. On the basis of his detailed studies of antipodal Hepaticae, Schuster (1963)
suggested it to be placed in a separate order Monocleales, including a single family Monocleaceae and
a single genus Monoclea. Schuster (1963), thus later recognised six orders amongst Hepaticae, viz.
(i) Calobryales, (ii) Jungermanniales, (iii) Metzgeriales, (iv) Sphaerocarpales, (v) Marchantiales, and
(vi) Monocleales.
Takakia lepidozioides, a new bryophytic genus discovered from Japan and Canada around the sixties
of the last century has been placed in a new order Takakiales with a single unigeneric family Takakiaceae
by Hattori and Inoue (1958) on the basis of some unique characters like (i) absence of rhizoids, (ii) very
low chromosome number (n = 4), and (iii) each leaf unit bearing two solidly parenchymatous structures.
Class Hepaticopsida thus includes seven orders: (i) Takakiales, (ii) Calobryales, (iii) Jungermanniales,
(iv) Metzgeriales, (v) Sphaerocarpales, (vi) Monocleales, and (vii) Marchantiales. Sandra Holmes
(1986) also divided the class Hepaticopsida into the same seven orders. An outline of her proposed
classification of Bryophyta is given in Article 2.1 of Chapter 2.
On the basis of their detailed study of sex organs and sporophytes of Takakia, Smith and Davidson
(1993) have shown the need to transfer Takakia from liverworts to mosses. Rashid (1998) has also
shown “the need to reclassify this genus to mosses”. However, some detailed studies are still needed and
till then this new unigeneric order created by Hattori and Inoue (1958), and Hattori and Mizutani(1958)
has been discussed here in this text as an order of the class Hepaticopsida.
Recent studies of molecular biology, genosystematics, phylogeny, diversification and classification
of bryophytes (Newton et al., 2000; Norris, 2003; Shaw and Renzaglia, 2004; Zander, 2006; He-
Nygren et al., 2006; and Troitskey et al., 2007) treat all bryophytes under an independent subkingdom
(Bryobiotina) and divided it further into three phyla, of which one is Marchantiophyta (= Liverworts
or Hepaticopsida or Hepaticae). For details, see Article 2.3.1, Chapter 2.
TAKAKIALES 3.5
3.5.1 What is Takakiales?
Takakiales is an interesting monogeneric order represented by only one genus, Takakia. It resembles
Calobryales of Hepaticopsida on one hand and mosses (Bryopsida) on the other hand. Smith and
Davidson (1993) studied the structure of its capsule and sex organs and suggested it to belong to mosses,
whereas in general, the organisation of its body structure and low haploid number of its chromosome
(n = 4) suggest that it belongs to Hepaticopsida. Detailed studies are, however, still needed to finally
decide about its inclusion either in Hepaticopsida or in mosses.
3.5.2 General Characteris cs

Some of the general characteristics of Takakiales are listed below:
1. Rhizoids are absent, and the function of absorption is taken up by the underground rhizomatous
stem.
2. A number of multicellular, usually unbranched, flask-shaped, mucilage-producing cells are
present on the underground rhizome.
3. Several thick, fleshy, green, cylindrical structures are present on the aerial portion of the plant.
These are called phyllids.
4. Phyllids are morphologically uniform and present in three rows. These are 3–5 cells thick
structures, each attaining a length of about 1 mm.
5. Plants apparently lack dorsiventrality.
6. Asexual reproduction is not well-known.
7. The gametophytes are heterothallic. The male plants have not been clearly reported.
8. The archegonia occur either singly or in groups of two or three on the female plants.
9. A small massive neck is made of six rows of neck cells. A fleshy venter is also present in the
archegonium.
10. The haploid chromosome number is n = 4.
3.5.3 Classifica on
Takakiales includes a single family (Takakiaceae) represented by only a single genus, Takakia.
Some required details of Takakia are discussed below.
TAKAKIA 3.6
3.6.1 Distribu on and Habitat
Takakia, the only genus of the family Takakiaceae of the order Takakiales, is represented by only two
species, viz. T. lepidozioides and T. ceratophylla. The name has been given after Takakia, a Japanese
botanist.
Takakia lepidozioides was discovered first by Hattori and Inoue in 1958 from the alpine zones of
Japan at an altitude of 2400–2800 metres. Later, it was also reported from British Columbia, Canada
and North Borneo at an altitude of about 3000 metres. Its gametophytes prefer to grow in moist shady
places on rocks, crevices and humus.
Takakia ceratophylla has been reported by Grolle (1963) from Sikkim in the eastern Himalayas in
India at an altitude of about 3800 metres. Later on, Hattori et al. reported this species from Aleutian
Islands of British Columbia in 1968. It grows on moist soil, ditches and also on the banks of streams.
3.6.2 Plant Body

The plant body is differentiated into a leafy shoot or gametophore and rhizome. The leafy shoot is
an aerial, erect, radially symmetrical structure attaining a length of about 1 cm. The rhizome is a
subterranean, branched and cylindrical structure (Fig. 3.1A,B). Rhizoids are absent. A large number of
mucilage hairs are present on both the leafy shoot as the well as on the rhizome.
26 � Bryophyta
Fig. 3.1 A-I, Takakia. A, Gametophores of T. lepidozioides; B, Gametophore of T. ceratophylla; C-E, Two,
three and four leafy segments of phyllids; F, A phyllid in cross sec on; G, A phyllid of complex
construc on in cross sec on; H, TS aerial shoot; I, Stalked archegonium.
Mucilage hairs are of two types. These are filamentous “closed” type and beaked “open” type
(Proskauer, 1962). The filamentous closed type of mucilage hairs are found on leafy shoots. These are
unbranched filaments, of which the terminal cell secretes mucilage through the unruptured cell wall.
The beaked open-type of mucilage hairs are found on both leafy shoots as well as on rhizomes. These
are usually branched and occur in clusters. Their terminal cell becomes beak-shaped and their tip bursts
and secretes mucilage.
Rhizome is almost colourless. It generally grows downwards and possesses no leafy structures.
There are no root caps. In dry conditions, the rhizome gets covered by mucilage. Rhizome has been
termed “stolon” by Schuster (1966), “caulid” by Hattori and Mizutani (1958), “rhizome” by Berrie
(1962) and also “root” by Grubb (1970).
Phyllids are the deep cylindrical structures present on the gametophores. They are isophyllous, i.e.
all alike and attain a length of about 1 mm. They are arranged triseriately on all sides of the leafy shoot.
Each phyllid is either bifid, trifid or quadrifid i.e. divided into two, three or four segments.
3.6.3 Anatomy
Anatomically, each leaf segment is divided into two parenchymatous cylindrical structures, each made
of a central row (Fig 3.1 F) or several rows (Fig 3.1G) of cells surrounded by a layer of epidermal
cells.
In a cross section, the aerial shoot (Fig 3.1H) is a somewhat elliptical structure and exhibits no evidence
of any dorsiventrality. Its cortex consists of one or two layers of thick-walled cells surrounding a few
centrally located thin-walled cells. The cells of the central strand, when mature, have no protoplasmic
content. Their end walls are perforated by many small pores. Hebant (1972) confirmed the conducting
role of this strand.
3.6.4 Apical Growth

The apical growth takes place by a single tetrahedral apical cell with three cutting faces. In this aspect,
Takakia resembles Calobryales and almost all Bryopsida.
3.6.5 Reproduc on
Asexual reproduction has not been reported in Takakia. With reference to sexual reproduction, Takakia
is a dioecious or heterothallic plant. However, male plants have also not been clearly reported so far in
this genus. Female plants bear archegonia. Either a single archegonium is produced at the apex of the
main shoot or two or three archegonia occur in groups.
The archegonium (Fig. 3.1 I) does not possess any protective structures around it, and it is thus
naked. It is a large, flask-shaped body mounted on a stalk. The large and massive nature of archegonium
shows its resemblance more with mosses than with Hepaticopsida. The neck is made up of six rows of
neck cells (Inoue, 1961). But, according to Hattori and Mizutani (1958), it consists of only four vertical
rows of cells. The venter is fleshy in nature. The jacket of the venter is usually two-layered.
Sporophytes have been discovered in Takakia by Smith and Davidson (1993). According to them,
the available details of the structure of sex organs and capsule bring Takakia close to mosses.
3.6.6 Affini es and Phylogene c Importance of Takakia

Takakia possesses several primitive features of phylogenetic significance, and on the basis of such
features, this genus is considered nearest to the ancestors of Hepaticopsida. Some such primitive
features are listed below:
1. Lowest haploid chromosome number in Hepaticopsida (n = 4).
2. Radial organisation of the plant axis with no differentiation into typical type of leaves.
28 � Bryophyta
3. Isophyllus, uniseriate to triseriate phyllids.

4. Mucilage hairs present on both rhizome and axis.
5. Presence of some thin-walled, water-conducting cells in the axis.
6. Complete absence of rhizoids.
7. Presence of slime papillae.
8. Presence of well-developed, massive archegonia having a primitive type of structure.
9. Particular lack of drought resistance.
All the above-mentioned features indicate that Takakia is a highly primitive genus and nearest or
quite close to the ancestral stock of liverworts. Presence of low haploid chromosome number (n =
4) suggests that Takakia represents relics of a race that appears to have died out. Only due to such
primitive features, Mehra (1969) opined that “Takakia is a living fossil in Hepaticae”.
Takakia resembles Calobryales in several characteristics including
1. Lack of rhizoids
2. Function of absorption taken up by underground rhizomatous stem
3. Presence of cluster of multicellular, mucilage-producing, flask-shaped cells on the rhizome
4. Isomorphous phyllids present in three rows
5. Radially organised shoots
6. Massive and primitive type of archegonial structure
7. Apparent lack of dorsiventrality
3.6.7 Is Takakia a Moss?

Smith and Davidson (1993) discovered sporophytes in Takakia. On the basis of their detailed studies
of its capsule and sex organs, they have proposed to reclassify Takakia along with mosses. They have
strongly pleaded that Takakia should be transferred from liverworts (Hepaticopsida) to mosses.
3.6.8 Takakia and Evolu on of Liverworts

Schuster (1966) opined that the low chromosome number (n = 4) in the gametophyte of Takakia
shows its resemblance with the lowest basic chromosome number of Chlorophyceae (green algae),
as in Ulothrix. In over 75% of all the liverwort species investigated so far, the basic haploid number,
however, is n = 9. This throws some light on the evolution of liverworts from algae.
3.6.9 Takakia: A Landmark in the History of Bryology

RM Schuster (1958), a noted American bryologist, worked on the relationships of Takakia with other
bryophytes. He may be quoted as “I am still not sure, but I suspect that a class parallel with Musci,
Hepaticae and Anthocerotae is at hand”. EV Watson (1964) elaborated it further that “If this were so,
the discovery of Takakia would be a landmark in the history of bryology”.
CALOBRYALES 3.7
3.7.1 What is Calobryales?

Calobryales is a small order of Hepaticopsida represented by only two genera, viz. Calobryum and
Haplomitrium. Its members are commonly described as “mosslike hepatics” because their radially
symmetrical gametophytes are erect leafy structures, in which the leaflike bodies are almost all alike
and remain arranged in three vertical rows. Calobryum is acrogynous because its apical cell develops
into an archegonium. Haplomitrium is, however, anacrogynous because there is no involvement of
apical cell in the development of archegonia. Schuster (1963) opined that separation of these two
genera on the basis of acrogynous or anacrogynous characteristics is unwarranted, and he thus united
and treated them as one single genus, namely Haplomitrium. A majority of bryologists, however, treat
them as two independent genera.

1. A branched, creeping, rhizome-like underground stem is present at the basal part of the
gametophytic plant body.
2. Erect leafy shoots develop from the underground rhizome-like stem.
3. Rhizoids are absent.
4. The leaflike structures are flat, unlobed, simple and entire.
5. The leaves are almost all-alike in shape and size. Out of three ranks of leaves, one is smaller
than the other two. This smaller one has been equated with the amphigastria of the order
Jungermanniales.
6. Plants are dioecious.
7. Female plants are acrogynous.
8. Arrangement of antheridia on the gametophore is different in both the genera of Calobryales.
In Haplomitrium, the antheridia arise in the axils of leaves while in Calobryum they are
present in groups on a terminal receptacle. Enlarged leaves form a cuplike structure around
the terminal receptacle.
9. Mode of development of sex organs is of the most primitive type than any other group of
bryophytes.
10. A massive calyptra, present in the sporophyte, is a peculiarity.
Calobryales includes only one family, Calobryaceae, having only two genera, viz. Calobryum and
Haplomitrium.
Calobryum, along with a few details of Haplomitrium, is discussed here in some detail.
CALOBRYUM AND HAPLOMITRIUM 3.8

Eight species of Calobryum have been reported so far while Haplomitrium is represented by only one
species (H. hookeri). Of the eight species of Calobryum, three have been reported from India. These
are C.blumii from Jawaii (Assam), C. indica from Darjeeling (West Bengal) and C. denundatum from
Himalayan regions. The remaining five species are C. adinum from Peru (Equador), C. gibboiae from
New Zealand, C. giganteum from Philippines, C. intermedium from Australia and C.mnioides from
Japan. Ram Udar, a noted Indian bryologist, reported Haplomitrium hookeri from Jawail (Assam)
30 � Bryophyta
and Vyas Shikhar (Western Himalayas). Haplomitrium also occurs in northern Europe and USA.
Calobryum chiefly occurs in tropical regions.
Both the genera of Calobryales occur in mesophytic conditions on forest floors, steepy clay banks
and other such surroundings in extremely shady and wet conditions.
3.8.2 Plant Body

Plant body of both Calobryum and Haplomitrium (Fig. 3.2 A,B) is gametophytic, and the gametophytes
contain a prostrate part and an erect succulent system. The prostrate underground part represents a
rhizomatous system from which develop erect and bright green leafy branches.
Fig. 3.2 A, Male gametophyte of Calobrium blumei; B, Sporophyte-bearing gametophyte

of Haplomitrium; C, TS stem
The leaves are arranged in three vertical rows on the stem. They possess almost the same shape and
size. Sometimes the leaves of one row are slightly smaller than those of two other rows. The leaves are
elliptical, contain the entire margin and attain a width of 2 to 5 mm. Sometimes they reach up to 10 mm
in Haplomitrium. Usually, the leaves are unistratose.
The underground part of the gametophytes is branched, extends downward and lacks leaves. Both
prostrate as well as erect systems lack rhizoids.
Anatomically, the stem (Fig 3.2C) contains enlarged cells in the outer zone. The cells of this zone
possess a few chloroplasts and many oil droplets. Narrow cells are present in the central zone. Its cells
are devoid of chloroplasts and contain only a few oil droplets. The cells of the central zone function as
a conduction system and resemble the hydroids of mosses.
Apical growth takes place by a pyramidal apical cell with three cutting faces.
3.8.3 Reproduc on
Sex organs in both Calobryum and Haplomitrium develop at the tip of the leafy branches, and both are
dioecious. Antheridia in male plants are surrounded by large-sized, rosette-forming leaves, giving an
appearance of mosses.
At the time of the development of antheridium, the antheridial initial divides transversely to form
a basal cell and an outer cell. The basal cell remains embedded in the gametophytic tissue while the
outer cell projects out and divides to form a primary stalk cell and primary antheridial cell. The primary
antheridial cell divides by three successive vertical divisions forming three jacket initials, surrounding
a single primary androgonial cell. Transverse and longitudinal divisions take place in the androgonial
cell and thus androcyte mother cells are resulted. These divide and form spermatozoids. The stalk cell
divides to form a stalk of the antheridium made up of several superimposed tiers, each made up of four
cells. A mature antheridium, thus developed, is a spherical body with a prominent stalk. Many such
antheridia surrounded by leaves are present near the shoot apex (Fig. 3.3A). The leaves form a cuplike
rosette. The antheridia are lateral and axillary in position.
Calobryum is acrogynous (in which terminal growth of the plants comes to an end due to utilization
of the apical cell in the formation of an archegonium), while Haplomitrium is anacrogynous (in which
archegonia are formed without the utilization of the apical cell). In Calobryum, the apical cell cuts off
segments to form about six or more archegonia, and ultimately the apical cell starts functioning as an
archegonial intital. Similar to antheridia, the archegonia are also lateral and axillary in position.
At the time of the development of archegonium, the primary archegonial initial forms a primary
axial cell surrounded by three jacket intitals. The primary axial cell functions directly as a central cell
without forming a primary cover cell. The archegonial neck is made up of only four rows of cells
because amongst the jacket cells only one divides. A mature archegonium contains a long neck
surrounding a vertical row of 16–20 neck canal cells. The venter has a breadth like that of a neck, and
at maturity it is only two cells thick. Usually, only one archegonium gets fertilised on an erect shoot.
3.8.4 Post-Fer liza on Changes

The diploid zygote divides by irregular divisions to form a multicellular embryo. Its lower or hypobasal
part develops into a haustorium and the upper or epibasal part develops into foot, seta, and the capsule.
Elaterophore is absent. The sporogenous tissue of the capsule is differentiated into elaters and spore
mother cells. Several spore tetrads are seen in the early stages. The developing sporophyte remains
surrounded by a massive calyptra.
A mature sporophyte (Fig. 3.3 B-D) bears an acuminate foot, long seta and a well-developed
capsule. Spore tetrads and several long elaters, tapering at both ends (Fig. 3.3 E), are present in the
capsule. Elaters bear helical thickenings. The capsule wall is one-celled thick throughout, except in
the apical regions (Fig. 3.3D). The jacket of the capsule possesses ornamentation showing unique
longitudinal bands, perpendicular to the capsular surface.
Seta becomes very long (8–30 mm) before dehiscence of capsule. The capsule comes out of the
calyptra. It dries, shrinks in length and dehisces, thus releasing the spores and elaters. Seta disintegrates
and collapses in the dehisced capsule. Spores are unicellular, green and possess chlorophyll. Each
spore germinates by forming a globose mass of cells. From the latter, the reclining branches come out
to form the primary rhizomatous system. Erect branches develop from this rhizomatous system. Thus
develops a young gametophyte bearing leaves on the erect branches.
32 � Bryophyta
Apical region of capsule
Spore tetrads
Elaters
Capsule wall
(one-celled thick)
An elater
Fig. 3.3 A, Ver cal sec on through the male apex of Calobryum blumei; B, Female plant with a mature
sporophyte; C-D, Capsule and its upper part enlarged showing spore tetrads, elaters and capsular
wall; E, An elater.
1. What are Hepa copsida commonly called?

2. What are Hepa copsida? Describe in about 50 words.
3. In recent years, some taxonomists have started calling Hepa copsida as _________.
4. Why do the members of Hepa copsida commonly called liverworts?
5. Enumerate at least 10 characteris c points of Hepa copsida.
6. Write a scien fic note on the classifica on of Hepa copsida in about 250 words.
7. What are Takakiales?
8. How many genera do Takakiales include?
9. Write any five dis nguishing points of Takakiales.
10. Give an illustrated account of some details of the life history of Takakia in about 500 words.
11. Where is Takakia found in India?
12. Describe affini es and phylogene c importance of Takakia.
13. Is Takakia a moss? Comment.
14. “Takakia is a landmark in the history of bryology”. Describe in about 50 words..
15. What are Calobryales? Write at least five of their characteris c features.
16. Calobryales is represented by only two genera, viz. Calobryum and ________ .
17. Describe the distribu on of Calobryales in about 50 words.
18. Write an account of some life-history details of Calobryum and Haplomitrium.
4
Hepaticopsida
(Jungermanniales)
Represented by about 250 genera and more than 9000 species, Jungermanniales represent the largest
order of Hepaticopsida. They are worldwide in their distribution and show the following general
characteristic features:
1. Plant body is gametophytic, and gametophytes are either thalloid or foliose.
2. Thalloid members contain a simple thallus while the foliose members have a leafy axis with
least differentiation of histological tissues.
3. Plant body is attached to any suitable substratum by rhizoids, which are always smooth-
walled.
4. Scales are generally absent.
5. Sex organs are represented by antheridia and archegonia, and plants are both monoecious
(e.g. Pellia epiphylla) as well as dioecious (e.g. Pellia neesiana). The monoecious species are
generally protandrous.
6. The antheridia are globose or subglobose.
7. In the process of antheridium development, “the primary antheridial cell does not undergo two
centric vertical divisions at right angles to each other, with the result that a quadrant of four
daughter cells is not formed” (Parihar, 1987).
8. Archegonial neck is almost as broad as the venter.
9. Archegonial jacket is generally made up of five vertical rows of cells.
10. The sporogonium is divisible into foot, seta and capsule.
11. Foot is somewhat bulbous, seta is highly elongated, and the capsule is globose or round and
bears spores and elaters.
12. The capsule is multistratose, i.e. its jacket is made up of two or more layers of cells in
thickness.
13. Prior to the first nuclear division, the spore mother cells generally become deeply four-lobed,
indicating the position of four spores in each of them.
14. The archesporium ultimately develops into fertile spores and sterile elaters.
15. The capsule generally dehisces by four valves on maturity.
Hepaticopsida (Jungermanniales) � 35
CLASSIFICATION 4.2
Classification of Jungermanniales is still in a very controversial stage, and different bryologists
include very different families and different number of genera in this taxa. Verdoorn (1932) divided
Jungermanniales in two artificial groups and treated them as two independent orders as under:
1. Jungermanniales Anacrogynae
2. Jungermanniales Acrogynae
Anacrogynae members are those in which archegonia develop on the dorsal surface of the prostrate
shoot, and there is no involvement of the apical cells in the formation of archegonia. On the other hand,
Acrogynae members are those in which archegonia develop at the apex of the shoot, and apical cell is
utilised in the formation of the archegonium.
Evans (1939) divided all members of the order Jungermanniales into three sub-orders as under:
1. Haplomitrineae
2. Metzgerineae
3. Jungermannineae
Sub-order Metzgerineae is equivalent to Jungermanniales Anacrogynae while the sub-order
Jungermannineae is equivalent to Jungermanniales Acrogynae of Verdoorn. Campbell (1936), another
well-known bryologist, opined that the sub-order Haplomitrineae of Evans has no resemblance with
Jungermanniales and should be treated under the order Calobryales.
Parihar (1987) followed Campbell (1936) and treated all members of the order Jungermanniales in
two sub-orders as under:
Sub-order 1: Jungermannineae (Jungermanniales Acrogynae)
Sub-order 2: Metzgerineae (Jungermanniales Anacrogynae)
The same classification has been followed in this text.
SUB ORDER JUNGERMANNINEAE

JUNGERMANNIALES ACROGYNAE 4.3
General Characteris cs
More than 75% of the members of Hepaticopsida belong to the sub-order Jungermannineae of the
order Jungermanniales. There are about 220 genera and more than 8500 species in this sub-order. Its
members show the following general characteristics:
1. The gametophytic plant body is usually differentiated into stem and leaves, and plants remain
attached to the substratum by a large number of rhizoids.
2. A very definite type of segmentation process is initiated by an apical cell, which is pyramidal
with three cutting faces.
3. Three rows of leaves are usually present on the stem or axis. Of these three rows, two rows of
leaves are laterally placed and made up of leaves of normal size while the third row of leaves
is present on the lower side, are smaller-sized and called amphigastria. In some genera, the
amphigastria are either highly reduced or even absent.
4. Dichotomous branching of the axis is never present.
5. The antheridia develop, either singly or in small groups, in the axils of modified leaves.
36 � Bryophyta
6. The archegonia develop at the apices of the stem or branches.

7. The apical cell is utilised in the formation of the last-formed archegonium. Due to this, the
vegetative growth of the axis or a branch stops after the formation of the last archegonium.
8. Fertilization usually takes place through the agency of water, present in the capillary spaces
existing between the plants and overlapping leaf surfaces.
9. The sporogonia are terminal on the branches or axis. A mature sporogonium usually consists
of an indistinct foot, short seta and a globose capsule.
The sub-order Jungermannineae contains 17 families, of which only two (Porellaceae and
Frullaniaceae) are briefly discussed here with some life-history details of Porella of Porellaceae and
Frullania of Frullaniaceae.
PORELLACEAE 4.4
Porellaceae is also known by the name Madothecaceae. It shows the following major features:
1. The rhizoids form tufts at the bases of the amphigastria.
2. The leaves show incubous arrangement, i.e. if seen from the above, the forward edge of each
leaf overlaps the hind edge of the leaf immediately next above to it on the same side.
3. The lobule or postical lobe of the leaf is quite distinct.
4. The amphigastria or underleaves are large-sized.
5. The postical lobes or lobules are replaced by lateral branches.
6. The perianth is bilabiate, well-developed, inflated, and contains almost compressed mouth.
7. At the time of dehiscence, the valves split about halfway down the capsule.
The life history of Porella is discussed here.
PORELLA 4.5
4.5.1 Systema c Posi on
Division—Bryophyta
Class—Hepaticopsida
Subclass—Jungermanniae
Order—Jungermanniales
Sub-order—Jungermannineae (Jungermanniales acrogynae)
Family—Porellaceae (=Madothecaceae)
Genus—Porella L. (=Madotheca Dum.)

Over 180 species of Porella have so far been reported, of which most occur in tropical regions and
only a few species are found in temperate regions of the world. About 35 species have been reported
from India, mostly in the Himalayan regions and a few also in south India. Plants grow on moist shady
rocks, stones and also on the bark of the trees. A few species grow on moist soil. P. platyphylla, a robust
liverwort (Watson, 1967), occurring commonly in limestone regions of Europe, North America and
several Asian countries, is the most widely distributed species.
Some of the common Indian species along with the regions of their common occurrence are Porella
gollani (Mussoorie, Garhwal), P. plumosa (Mussoorie, Chamba, Dalhousie), P. variabilis (Mussoorie),
P. macroloba (Kumaon, Kullu valley), P. angusta (Simla, Kashmir), P. hastata (Mussoorie), P.
densiramea (Chamba), P. densifolia (Kumaon) and P. boreilli (Kashmir).
GAMETOPHYTIC PHASE
4.5.3 External Morphology of Gametophyte
The plant body is gametophytic, and the gametophytes are large, flat, compact, dorsiventral, well-
branched and foliose structures, divisible into rhizoids, axis and leaves (Fig. 4.1 A-C). The plants reach
up to 15 cm or more in length.
The axis or stem is prostrate, dorsiventrally flattened and bi- or tri-pinnately branched. Each branch
of the axis remains covered by three rows of leaves, of which two rows of larger leaves are present
on the dorsal side, while the third row of smaller leaves is present on the ventral side of the axis. The
smaller leaves are called amphigastria. The larger leaves of the dorsal side of the axis are bilobed,
and the lobes are unequal-sized. The upper, oval-shaped lobe of each larger leaf is large and called
the antical lobe, while the lower smaller lobe is called the postical lobe. The postical lobe contains
Dorsal leaf Lobe
Lobule
Stem
Ventral
leaf
Megaphyll
Rhizoid
B
A
Microphyll
Fig. 4.1 A-C, External features of the gametophyte of Porella. A, Dorsal view of a branch of P. platycarpa;
B, Ventral view of a branch of P. platycarpa; C, A part of the gametophy c plant body of P. gollani
38 � Bryophyta
an acute apex and appears like a separate leaf. The posterior margin of the leaf underlines the anterior
margin of the older leaf and this arrangement is called incubous type. The amphigastria resemble the
postical lobes of the larger leaves.
A large number of the smooth-walled rhizoids arise on the lower side of the stem. The plants remain
closely pressed to the moist substratum, and the absorption of water takes place directly through the
cells of stem and leaves. The main function of rhizoids is thus the anchorage of the plants to the
substratum.
4.5.4 Anatomy of Axis

There is little or no differentiation of tissues in the young axis. Old stems show some differentiation
of outer cortical cells and central medullary cells. Usually the cells of the cortex are small and thick-
walled while that of the medulla are large and thin-walled. Almost all cells are parenchymatous.
4.5.5 Anatomy of Leaf

Internally, the leaves are very simple in structure, consisting of a single layer of isodiametric, polygonal
cells, each containing abundant chloroplasts. There is no midrib. Each cell in P. acutifolia contains
7–19 oil bodies (Udar, 1971).
4.5.6 Apical Growth

It takes place by the activity of a tetrahedral apical cell with three cutting faces. The apical cell is
pyramidal in shape, and it regularly cuts three sets of segments, of which two sets are dorsal and lateral
while the third set is ventral. Each segment develops into a leaf and a part of the stem.
4.5.7 Vegeta ve Reproduc on

Plants have been reported to reproduce vegetatively by fragmentation, especially when they grow under
humid conditions. Schiffner described the presence of discoid gemmae as the means of vegetative
reproduction in Porella rotundifolia growing in Brazil, but this report has been denied by Degenkolbe
(1938). No other authentic report of vegetative reproduction is available in Porella.
4.5.8 Sexual Reproduc on

The sexual reproduction is oogamous. Porella is dioecious. The male plants are smaller in size than the
females. The antheridial branches arise more or less at right angles to the main axis (Fig. 4.2). Usually,
the female plants are larger and bear the archegonial branches which are shorter than that of antheridial
branches.
4.5.9 The Male or Antheridial Branch

Usually, the antheridial branches arise at right angles to the main axis and have more compactly arranged
leaves than the vegetative branches. The leaves of the antheridial branches are closely imbricated and
called ‘bracts’. A single antheridium develops in the axil of each bract of the male branch.
Fig. 4.2 A part of the leafy shoot of Porella platycarpa having one antheridial branch in ver cal view
1. Development of Antheridium
The development of antheridium in Porella resembles with Pellia and other members of Jungermanniales,
and a general account of the same is given below:
A superficial dorsal cell starts to function as an antheridial initial. It becomes papillate and divides
by a transverse wall into an outer cell and a basal cell (Fig. 4.3 A,B). The outer cell remains protruded
above the general surface of the thallus while the basal cell usually remains embedded in the thallus
tissue. The outer cell divides transversely into an upper primary antheridial cell and a lower primary
stalk cell (Fig. 4.3C).
The stalk of the antheridium develops from the primary stalk cell. The primary antheridial cell
divides by a median vertical division (Fig. 4.3D) to form two equal-sized daughter cells. Each of these
daughter cells divides by a somewhat periclinal division forming two unequal cells (Fig. 4.3F). The
smaller of these two unequal cells represents the first-jacket initial while the larger one divides by
another periclinal division to form an outer second-jacket initial and an inner primary androgonial
cell (Fig. 4.3E). Two triangular, centrally-located primary androgonial cells surrounded by four
jacket initials are seen in the cross section of the young antheridium at this stage (Fig. 4.3G). Both the
primary androgonial cells divide by repeated transverse and vertical divisions to form a large number
to androgonial cells (Fig. 4.3 H-J), of which the last generation is of the androcyte mother cells.
Repeated anticlinal divisions in the jacket initials give rise to a single-layered sterile jacket. Each
androcyte mother cell divides diagonally to form two androcytes, of which each metamorphoses into a
uninucleate and biflagellate antherozoid.
2. Mature Antheridium
The mature antheridium has a globular body and a long stalk (Fig. 4.4). The stalk consists of two
rows of cells while the body remains surrounded by a sterile jacket. The jacket is single-layered in
the upper part while it is bi- to tri-layered at the base. The jacket encloses the androcytes, which soon
metamorphose into biflagellate and coiled antherozoids.
40 � Bryophyta
Antheridial initial
Outer Primary
cell antheridial
cell
Basal Primary
cell stalk
cell
A
B
Primary androgonial cells
1st Jacket initial

2
2 3
1 1 1 1
2 3 2
F
E 2nd jacket
D G
initial
Androgonial Jacket
cells
Jacket
Stalk
I
H J
Fig. 4.3 A-J Development of antheridium in Porella and other Jungermanniales
Fig. 4.4 A part of an antheridial branch of Porella bearing antheridia in the axil of leaves
3. Dehiscence of Antheridium
One-celled-thick distal region of the jacket of the antheridium is thinner, and when it is near to the
water, it bursts open into a number of irregular lobes. These lobes curl back strongly. Because of this,
the entire mass of androcytes is forced out into the water. This mass of androcytes along with several
antherozoids soon ruptures, and thus the antherozoids are liberated.
4.5.10 The Female or Archegonial Branch

The archegonial branches are smaller than the antheridial branches, and are hence much less distinct.
At the tip of these female branches are present the archegonia. The first two or three leaves in each
archegonial branch are sterile and do not bear archegonia. When young, the first few archegonia develop
in strict acropetal succession. However, ultimately the apical cell of the archegonial branch starts to
function as an archegonial initial. Any further growth of the archegonial branch is thus stopped.
1. Development of Archegonium
In the development of the archegonia also, Porella shows strict resemblance with Pellia and other
Jungermanniales, and a general account of the same is given below:
The development of archegonium starts from a single superficial cell, which soon becomes papillate
and starts to function as an archegonial initial (Fig. 4.5A). It first divides transversely into a lower
basal cell and an upper outer cell (Fig. 4.5B). The basal cell divides transversely into a stalk initial
and a basal cell (Fig. 4.5C). The outer cell divides by three intersecting vertical divisions, resulting
ultimately into a central primary axial cell surrounded by three peripheral initials (Fig. 4.5 D-F and
D1-F1). One of these three peripheral initials is small and does not divide by any vertical division while
the remaining two peripheral initials divide by vertical divisions. This results in the formation of five
jacket initials (Fig. 4.5 G1).
Each jacket initial divides transversely to form the lower venter initial and upper neck initial (Fig.
4.5G). The neck initials divide only transversely and form a neck consisting of five vertical rows of
cells. The venter initials divide several times to form the globular venter of the archegonium.
The primary axial cell (Fig. 4.5 E) divides by a transverse division to form an upper primary cover
cell and a lower central cell (Fig. 4.5F). Yet another transverse division in the central cell divides it
into an upper primary neck canal cell and a lower primary ventral cell. The primary neck canal
cell divides, redivides transversely and gives rise to 6–8 neck canal cells, while the primary ventral
cell divides transversely only once and gives rise to an upper ventral canal cell and a lower egg
(Fig. 4.5 I).
The primary cover cell (Fig. 4.5F) divides by two vertical divisions at right angles to one another to
form four cover cells (Fig. 4.5 G).
2. Mature Archegonium
The mature archegonium (Fig. 4.5I) contains a long neck and a slightly globular venter. The neck
consists of five vertical rows of cells and encloses 6 to 8 neck canal cells. The wall of the venter is bi-
layered and encloses a ventral canal cell and an egg.
42 � Bryophyta
Fig. 4.5 A-I Generalized process of archegonium development in Porella and other
Jungermanniales (Note D1-G1, which are the cross sec ons)
4.5.11 Fer liza on

The fertilization takes place with the help of water, present usually in the fine spaces existing between
the closely growing male and female plants or in between the overlapping leaf surfaces of two plants.
Male and female plants in Porella grow in very dense, compact patches. The antherozoids from the
antheridia reach up to the archegonia of female plants with the help of water. The two gametes fuse and
form a diploid zygote.
SPOROPHYTIC PHASE
4.5.12 Development of the Sporogonium
The zygote starts to increase in size and almost fills the entire cavity of the venter. Soon, it divides
transversely into an upper epibasal cell and a lower hypobasal cell (Fig. 4.6 A,B). One more transverse
division in the epibasal cell makes the three-celled stage of the young embryo (Fig. 4.6 C). There is no
further division in the hypobasal cell and it functions as a suspensor (Fig. 4.6D, E) or haustorium. The
derivatives of the epibasal cell are responsible for the formation of the whole sporogonium. The upper
two cells of the three-celled embryo divide first by transverse and then by vertical divisions (Fig. 4.6D,
E) to form a multicellular structure, in which it is very difficult to draw a demarcation line between the
capsule and the seta.
Fig. 4.6 A-F Development of the sporogonium in Porella bolanderi
The terminal segments of this multicellular embryo divide by a periclinal division to form an outer
layer of amphithecium and an inner tissue of endothecium (Fig. 4.6F). The amphithecium divides by a
few periclinal divisions to form a 3- to 4-layered jacket of the capsule. The archesporium is endothecial
in origin in Porella. The cells of the sporogenous tissue get arranged in distinct rows radiating from
the base of the capsule. Some of the sporogenous cells show no further division and start to function
as young spore mother cells, while other sporogenous cells elongate and develop into young elaters.
Soon, the spore mother cells become four-lobed. Their diploid nucleus divides reductionally to form
four haploid nuclei, of which one enters into each lobe of the spore mother cell. The infolding of the
walls of the four-lobed spore mother cell takes place and four haploid spores are formed from a spore
mother cell. The elaters are short, have tapering ends, and each contains one or two spiral thickenings
(Fig. 4.8 A).
The seta is present in between the capsule and the foot. It is short and gradually merges into the
basal part of the capsule.
The foot is present at the base of the young sporogonium. It is bulbous and grows downward into
the tissue of the female branch. A short tubelike structure, formed from the outer tissue of the stem of
the female branch, encloses the foot as well as the seta.
Three definite coverings or envelopes surround the sporogonium when it is young. These are
calyptra, perianth and involucre. The calyptra is multilayered and surrounds the capsule completely
44 � Bryophyta
in its young stages. The perianth develops by the fusion of two uppermost bracts. The enlarged bracts,
covering the base of the perianth, form the involucre.
4.5.13 Mature Sporogonium

The mature sporogonium (Fig. 4.7) consists of foot, seta and capsule. The foot is less conspicuous,
globular or oval and remains embedded in the tissue of the stem of the archegonial branch. The seta
is short and present in between the foot and the capsule. The capsule is surrounded by a 3- to 4-celled
thick wall, which encloses fertile spores and sterile elaters (Fig. 4.8). Three definite envelopes (calyptra,
perianth and involucre) also surround the sporogonium.
Fig. 4.7 L.S. of a mature sporogonium of Porella spp.
4.5.14 Dehiscence of Capsule and Dispersal of Spores

In mature sporogonium, the seta elongates, and because of this elongation, the capsule pushes and
ruptures the outer protective coverings of calyptra, perianth and involucre. The capsule opens by four
valves, and through this opening the spores are dispersed.
YOUNG GAMETOPHYTE
4.5.15 Spores
The spores are round, spherical or globular bodies varying from 0.03 to 0.05 mm in diameter. The
wall of each spore is bilayered, of which the outer layer is smooth, papillose or echinate and called
exospore, while the inner layer is smooth and called endospore. Sometimes, the remnants from the old
spore mother cells also get deposited on the exospore in the form of a third layer called outer exospore
or perinium. The spores are uninucleate.
4.5.16 Development of Young Gametophyte

The spores start dividing while they are still inside the undehisced capsule. Some of them become
multicellular even within the capsule wall. With the help of some transverse and vertical divisions in the
spore, a multicellular protonema is resulted (Fig. 4.8 B-F). Soon, an apical cell of the shoot becomes
differentiated in this multicellular protonema. A few rhizoids also develop at a slightly later stage.
Fig. 4.8 A-F, Porella platyphylla. A, An elater; B-E, Showing germina on of spore;
F, Mul cellular thalloid structure which develops into new gametophyte
FRULLANIACEAE 4.6
Frullaniaceae includes only three genera (Frullania, Jubula and Neohattoria), and some of the general
characteristics of this family are listed below:
1. Plants are usually large-sized and predominantly reddish brown or deep green in colour.
Sometimes they are purplish brown in colour.
46 � Bryophyta
2. The stem is usually prostrate and pinnately or bipinnately branched.

3. Three rows of leaves are present on the stem, of which two rows are of lateral leaves and one
row of underleaves called amphigastria.
4. Characteristic Frullania-type of branches are present in the majority of members. In this type,
a “branch develops from the ventral half of the lateral segment replacing the lobule of the leaf.
The lateral leaves are complicate-bilobed and each leaf is divided into a flat antical lobe and a
cylindrical saccate or postical lobe (lobule)” (Parihar, 1987).
5. Usually, the lobule or postical lobe of the leaf is divided into two parts, viz. stylus and lobule
proper.
6. The amphigastria or underleaves, are usually bilobed.
7. The tuft of rhizoids is present usually at the base or middle of the amphigastria.
8. Archegonia vary from 2 to 12 in a group.
Some life-history details of only Frullania are discussed here.
FRULLANIA 4.7
Predominantly a tropical genus, Frullania is represented by more than 700 species. Many of these
species also occur luxuriantly in subtropical regions of the world. More than 40 species of Frullania
have been reported from India. It grows commonly on wet grounds and generally on moist rocks as a
lithophyte. Plants are found to form extensive mats on tree trunks, rocks, etc.
Plant body (Fig. 4.9A) is gametophytic, and the gametophytes are reddish-brown or blackish in
colour. Each plant is differentiated into a well-branched prostrate, stemlike central axis possessing
many leaves. The axis is pinnately or bipinnately branched. Many smooth-walled, unicellular and
unbranched rhizoids are present in tufts at the base of the axis or middle of amphigastria (underleaves).
Rhizoids keep the thallus attached to the substratum. The leaves are attached on the axis or stem in three
rows i.e. two rows of lateral leaves and one row of amphigastria or underleaves. Each leaf contains
a large antical lobe and a small postical lobe or lobule. The antical lobe of the lateral leaf is the well-
expanded part. It is sub-orbicular or obliquely ovate. Its margin is entire. The postical lobe or lobule
is cucculate, galeate or saccate, and either remains open or forms a water sac. Stylus a short subulate
process, is present on the postical lobe. It is actually situated between the stem and the postical lobe.
The amphigastria or underleaves are round, notched or deeply-lobed and smaller than that of the lateral
leaves. The amphigastria, together with stylus, form capillary spaces which retain water. In the female
plants or branches, the archegonial group is surrounded by 2 to 5 pairs of perichaetial bracts, of which
the uppermost bracts are laterally fused to form the perianth. The small lobe of the leaves forms a water
sac or pitcher. The function of the water sacs is to hold a part of the rainwater, which would otherwise
be lost. The stylus, underleaves and lateral leaves also form capillary spaces which retain water.
The anatomy of the stem (Fig. 4.9D) exhibits very little cell specialisation or tissue differentiation.
It is circular in outline. The epidermis is not well-defined and the cortical and central medullary zones
can be recognised. The cortical region is peripherally located and its cells are smaller than that of the
centrally-located medullary zone. Cortical-region cells are pigmented in the older portion of the axis.
The medullary region cells are thin, colourless and quite large. Conducting tissue is absent.
Fig. 4.9 Frullania dilatata. A, A part of female gametophyte showing perianth; B, Ventral view of the shoot
with underleaves removed; C, A part of the shoot from below (F.squarrosa); D, Cross sec on of axis
Apical growth takes place by a tetrahedral apical cell with three cutting faces. Three sets of segments
are cut off from this apical cell, of which two sets are dorsolateral while the third set is ventral. From
each of these sets develop a leaf and a portion of stem subtending it. The young primordium of each
dorsolateral leaf divides into two parts. The upper part of this young leaf primordium develops faster,
bends over the growing point and forms the leaf lobe. The lower smaller half develops into a hollow
structure which matures or develops into a pitcher or water sac.
Vegetative reproduction takes place by (i) death and decay of the older parts, leaving the ordinary
branches to develop into new plants; and (ii) gemmae, which are the multicellular, nodular outgrowths,
which are discoid to irregular in outline and develop into new plants on being detached.
48 � Bryophyta
Fig. 4.10 Frullania dilatata A-B, Male plant bearing antheridial branches (A) and a male branch bearing
antheridia (B); C, A mature antheridium; D, LS perianth showing two archegonia; E, Part of the
plant showing perianth.
Sexual reproduction is oogamous. Both dioecious as well as monoecious species are present in
Frullania. Antheridia and archegonia develop on different branches in monoecious species.
Antheridia (Fig. 4.10 A-B) develop in the axils of perigonial bracts of which two or more pairs
develop on short lateral branches. These bracts are arranged in a densely intricate manner. Usually, two
antheridia develop in the axis of each bract.
The development of the antheridium follows the same pattern as that of Porella (Fig. 4.3 A-J) and
Jungermanniales and has been described in detail under Article 4.5.9(1). The mature antheridium is
a globose or spherical body (Fig. 4.10 C) with a slender stalk of two rows of cells. The globose body
of the antheridium is differentiated into a centrally-located mass of androgonial cells surrounded by a
sterile layer of jacket. Each androgonial cell divides diagonally into two androcytes. The protoplast of
each androcyte metamorphoses into a typically biflagellate antherozoid.
Clusters of archegonia (Fig. 4.10 D-E) occur at the apices of short lateral branches of the female
branch in monoecious species and female plant in dioecious species of Frullania. Each cluster contains
two to four archegonia (Fig. 4.10 D). The leaves associated with archegonia on the female branch are
called perichaetial bracts. Two to five pairs of perichaetial bracts surround each cluster of archegonia.
Each perichaetial bract is bilobed or dentate and larger than the foliage leaves. The bracts situated
in the upper part of the female branch are laterally fused and form the perianth (Fig. 4.10E). The
perianth is inversely heart-shaped. It remains contracted to form a tubular mouth. The development of
archegonium (Fig. 4.5 A-I) follows the same pattern as that of Porella and other Jungermanniales and
has been described in detail under Article 4.5.10(1). The mature archegonium is a flask-shaped body
containing a basal swollen venter and long narrow neck . A small egg cell and a ventral canal cell are
present in the region of the venter. Five vertical rows of neck cells form the neck. They surround an
axial row of as much as eight neck canal cells.
Fertilization takes place as usual with the help of water and follows the same pattern as described
for Porella (Article 4.5.11). The two gametes fuse and form a diploid zygote.
The diploid zygote divides and redivides to form a sporophyte. First, it divides by a transverse wall
forming an epibasal and a hypobasal cell (Fig. 4.11A). The epibasal cell divides by a transverse wall
and the hypobasal cell by a vertical wall to form a four-celled stage (Fig. 4.11B). Thus formed two
upper cells divide by two vertical walls at right angles to one another. Two lower cells (hypobasal cells)
divide by another vertical division at right angles to the first one. The so-formed young embryo now
consists of three tiers, each tier made up of four cells. Of these three tiers the upper tier develops into
capsule, the middle tier into seta, and the lowermost tier develops into the foot of the mature sporophyte.
Repeated irregular divisions (Fig. 4.11 C-E) take place in the lowermost tier which ultimately develops
into an absorptive organ called foot (Fig. 4.11E). Its cells grow into papillae. The cells of the middle
tier divide by longitudinal divisions to form seta. The cells of the uppermost tier are responsible for
the formation of capsule. They divide by periclinal division to form four outer or peripheral cells
representing amphithecium and four central or axial cells representing the endothecium.
The archesporium (Fig. 4.11 E,F) is endothecial in origin. The endothecium cells divide only
longitudinally to form as many as two hundred or more cells forming a mass of cells. These cells
differentiate into fertile sporocyte-producing cells and sterile elater-producing cells (Fig. 4.11H).
Sporocyte-producing cells form spore mother cells (Fig. 4.11I) by repeated transverse divisions. The
elater-producing cells remain undivided and grow into long and narrow elaters with a single broad
spiral band. Round or cubical spore mother cells and long elaters are thus present in regular alternate
bands (Fig. 4.11I) The capsule wall contains two layers which develop from the amphithecium. The
archegonial venter develops into calyptra which surrounds the young sporogonium. The perianth also
encloses the sporogonium along with the calyptra (Fig. 4.12A).
50 � Bryophyta
Fig. 4.11 A-I Showing stages of embryogeny and development of young sporophyte in Frullania dilatata.
Mature sporophyte of Frullania consists of a blunt foot, short seta and a globose capsule. The
foot remains embedded in the tissues of the female branch and functions as an absorbing organ. The
seta is very short and attains a length of only about 1mm. The mature seta is about 8 to 9-cells thick
and supports the capsule at its distal end. The capsule is globose and remains surrounded by a two-
layered wall. Their walls are sclerified. The cells of the outer layer contain rodlike fibres, particularly
at the corners of their lateral wall. The walls of the cells of the inner layer of capsule possess irregular
network of thickening fibres. The capsule wall encloses many spores and elaters. The elaters extend
from roof to the floor of the globular capsule. The elaters are flattened, nonspiral and each possesses
a trumpet-shaped lower end. The spores are round or oblong in shape. Each spore is uninucleate and
remains surrounded by a two-layered wall, of which the outer layer is rough or tuberculate and the
inner layer is generally smooth. Many chloroplasts are also present in the spore.
Fig. 4.12 Frullania dilatata. A, Young sporophyte; B-H, Various stages of the development of gametophyte.
At the time of dehiscence, the capsule wall splits suddenly into four valves. Spores are flicked away
in the air by the sudden movements of the contracting elaters.
Each spore germinates into a young gametophyte. Germination of spore starts while it is still inside
the capsule. The first division of the spore is transverse (Fig. 4.12 B-C) forming two cells, of which
the upper cell now divides by a vertical division (Fig. 4.12 D). Divisions in all the three planes now
take place, resulting into a young globose protonema of 50–60 cells (Fig. 4.12 E-F). Few rhizoids
now develop from the young multicellular sporeling (Fig. 4.12 G). Primary leaves followed by the
development of juvenile leaf and underleaf now start developing from the young gametophyte (Fig.
4.12 H).
52 � Bryophyta
SUB ORDER METZGERINEAE

JUNGERMANNIALES ANACROGYNAE 4.8
About 25 genera and more than 600 species constitute the sub-order Metzgerineae. Its members show
the following general characteristics:
1. Gametophytic plant body is usually thalloid, and rarely is differentiated into stem and leaves.
2. Usually, the gametophtes are dorsiventral and prostrate.
3. Being basically thalloid, with some also having leafy thalli, and rarely differentiated into stem
and leaves, members of Metzgerineae are commonly named multiform-thallose-hepatics.
4. Sex organs remain scattered on the dorsal surface of the thalloid plant body. But they are also
sometimes present on highly reduced specialised branches.
5. Apical cell cuts off young segments which develop into archegonia. Differing from
Jungermannineae, the apical cell in these members is not utilised in the formation of an
archegonium.
6. The fully developed sporophyte is dorsal in position, and it is located some distance behind the
apex of the plant body.
7. The wall of the capsule is not strictly two-layered. It is 2-5 layers thick.
Parihar (1987) followed Evans (1939) and mentioned that the sub-order Metzgerineae includes eight
families, viz. Treubiaceae, Fossombroniaceae, Pelliaceae, Blasiaceae, Pallaviniaceae, Metzgeriaceae,
Riccardiaceae and Monocleaceae. But, Rashid (1998) mentioned that Metzgeriales (multiple thallose
hepatics) include as many as 12 recognised families.
Only three genera, namely Pellia of Pelliaceae, Riccardia of Riccardiaceae and Fossombronia of
Fossombroniaceae are described in this text.
PELLIACEAE 4.9
Some characteristics of Pelliaceae are listed below:
1. Plant body is thalloid, often with lobed or sinuous margins.
2. Sex organs (antheridia and archegonia) remain scattered on the dorsal surface of the thalloid
plant body.
3. Capsule is a globular or oval body.
4. A basal elaterophore is present in the capsule.
PELLIA 4.10
Order—Jungermanniales
Sub-order—Metzgerineae (Jungermanniales Anacrogynae)

Family—Pelliaceae
Genus—Pellia

Pellia is a thalloid liverwort spread widely in north temperate regions of the world. It is represented
by four species, namely P. epiphylla, P. endiviaefolia, P. neesiana and P . columbiana. Jones (1958)
recognised a fifth species, P. borealis. Three of its species occur widely in the Himalayas, including P.
epiphylla and P. endiviaefolia according to Kashyap (1929). P. epiphylla is quite common in Sikkim,
Darjeeling and eastern Himalayas while P. endiviaefolia occurs commonly in western Himalayas and
Kumaon regions.
Parihar (1987) mentioned that it was Raddi who formed this genus and named it Pellia “in honour
of Leopoldo Pelli-Fabbroni, a lawyer of Florence, Italy, and a friend of Raddi”.
Pellia loves to grow in moist shady places, particularly by the sides of moist rocks, ditches, streams
and other similar surroundings. It also sometimes grows submerged under flowing water.
4.10.3 External Morphology of Gametophyte

Plant body is gametophytic, thalloid, prostrate, thin, dorsiventral and appears to be dichotomously
branched. Margins of thallus are sinuous (Fig. 4.13 A-C). A well-defined midrib is present on the
Fig. 4.13. A, Thallus of Pellia epiphylla; B, A male thallus; C, A female thallus; D-E, TS thallus;
F, Part of longitudinal sec on of thallus
54 � Bryophyta
dorsal surface of the thallus. The growing point is situated in a notch at the anterior end of each branch
of the thallus. Thousands of smooth-walled, unicellular rhizoids are present in the midrib region on the
ventral surface of the thallus. Tuberculate rhizoids and scales are absent.
The thallus becomes long, narrow, ribbonlike, thin and delicate, when growing either in the water
or in highly moist and shady places. In such conditions, the thallus also contains very thin margins and
a distinct midrib. The plant body becomes broader, robust and elongated on damp grounds. It becomes
shorter, thicker, stouter and stunted with no clear midrib when growing on dry sandy soil.
4.10.4 Anatomy of Thallus

Anatomically, the thallus is very simple with least differentiation of tissues and mainly consists of
parenchymatous cells (Fig. 4.13 D, E). The midrib region is many-layered, thick and remains projected
towards the lower side. Marginal sides or wings of the thallus are thin and only single-layered. Abundant
chloroplasts are present in the cells of the wings and upper layers of the midrib region. Cells of the lower
regions of the midrib contain only a few chloroplasts or are completely devoid of chloroplasts. Most of
the cells of the thallus contain starch grains. The thallus also contains some oil-filled cells. Some yellow
or brown interlacing thickenings form a network of bands in the cells of the median region in species
such as Pellia neesiana and Pellia epiphylla. These bands run vertically as well as transversely, and
are seen clearly in mature thalli of P. epiphylla (Fig. 4.13 F). These thickenings are, however, absent in
Pellia endiviaefolia. The thallus is bounded on both the sides by a layer of epidermis. Smooth-walled
rhizoids develop from some cells of the lower epidermis in the midrib region.
4.10.5 Apical Growth

In Pellia, the growth takes place by an apical cell, situated in the notch. In P. endiviaefolia, the apical
cell is cuneate or wedge-shaped with four cutting faces (Fig. 4.14 A), of which two are lateral, one
dorsal and one ventral. But in Pellia epiphylla, it (Fig. 4.14 B) is lenticular and has a posteriorly convex
and two lateral cutting faces. Mucilage-secreting glandular hairs are also present in the regions of the
growing points of the thallus. In P. epiphylla, the cells cut off from the posterior face, divide repeatedly
and form the midrib of the thallus, while the lateral segments divide repeatedly and form the wings.

Pellia reproduces vegetatively by (i) adventitious shoots, which may develop from the margins of the
dorsal surface of the thallus, and (ii) death and decay of the older portions of the thallus, as in many
other thalloid bryophytes.

Plants may be monoecious (Pellia epiphylla) or dioecious (P neesiana, P. endiviaefolia). Antheridia
develop first in monoecious species, i.e. they are protandrous, as P. epiphylla.
4.10.8 Antheridia and Their Development

The antheridia develop in two to four rows on the dorsal surface of the thallus along the midrib (Fig.
4.13B). They appear as wartlike outgrowths along the midrib of the thallus. Each such outgrowth is
actually the antheridial cavity containing a single antheridium. Each antheridial cavity opens by an
opening on the dorsal surface.
Fig. 4.14 Apical cell and its segmenta on in Pellia endiviaefolia (A) and P. epiphylla (B); C, A much enlarged
mature antheridium of Pellia epiphylla; D, A free antherozoid of P. epiphylla
Development of antheridium follows almost the same pattern as that of other Jungermanniales, and
has been described in detail for Porella under Article 4.5.9(1), Fig. 4.3 A-J.
The mature antheridium is almost a spherical structure borne in an antheridial chamber opening
outside by a small pore. It remains attached to the thallus by a short multicellular stalk, and its main
body remains surrounded by a single-layered antheridial jacket (Fig. 4.14 C), which encloses numerous
androcytes. Each androcyte produces a single antherozoid (Fig. 4.1D), which possesses a spirally
coiled body containing a nucleus. Two long flagella are attached at the anterior narrow end of the
antherozoids.
The antheridium dehisces when water finds its way into the antheridial chamber. The antheridial
apex ruptures, releasing the mucilaginous mass of cells. The released mass of cells spreads over the
water as a thin film, and thus dehiscence is completed. Spreading of the mass of cells over the water
surface in Pellia and other bryophytes is probably due to lowering of the surface tension. Androcytes
are thus carried up to the archegonial involucre. Walton (1943) opined that “androcytes reach the
archegonial involucre in 15 seconds in Pellia epiphylla”.
56 � Bryophyta
4.10.9 Archegonia and Their Development

Groups of 4 to 12 archegonia develop at the anterior end (Fig. 4.15A) of the thallus near the growing
point. A flaplike (Pellia epiphylla) or tubular (P. endiviaefolia) or cylindrical ( P. neesiana) involucre
protects the group of archegonia. The involucre develops from the cells of the thallus behind the
archegonial group. Short mucilaginous hairs accompany the archegonial group.
Development of archegonium follows almost the same pattern, as discussed earlier along with
Porella in Article 4.5.10(1) and depicted in Figure 4.5 A-I.
The mature archegonium (Fig. 4.15 B) is a shortly-stalked multicellular body with a dilated venter
and a long neck. Six to eight neck canal cells are enclosed in the neck which remains surrounded by
a jacket made up of five vertical rows of cells. The venter and lower part of the neck may become
bilayered prior to fertilization.
The process of fertilization and syngamy is similar to that of Riccia and discussed in detail under
Article 7.6.9. The ultimate result is the formation of a diploid zygote.
4.10.10 Sporophyte
The zygote increases in size and secretes a wall around itself. Cells of the venter grow actively and
form a well-developed calyptra, which keeps enclosing the young developing sporogonium for quite
some time.
Fig. 4.15 A, LS of thallus of Pellia epiphylla showing sex organs; B, A mature archegonium
Development of sporogonium starts first by a transverse division forming an upper epibasal and a
lower hypobasal cell (Fig. 4.16 A, B). There is no role of the hypobasal cell in the formation of proper
embryo. It simply forms a suspensor which functions as a haustorium. All major parts of the mature
sporogonium (such as foot, seta and capsule) are thus formed by the upper epibasal cell.
The epibasal cell divides first by a vertical wall followed by transverse division at right angles to the
first division, thus forming four cells. All these four cells now divide by vertical divisions, thus forming
two tiers of four cells each. The lower tier of four cells now divide and redivide to form the foot and
seta. The foot is well-developed and attains a conical shape. Its projecting edges grow upwards, overlap
the basal part of the seta and appear like a collar. The upper tier of four cells of the young embryo
divide periclinally to form the central endothecium and outer or peripheral amphithecium (Fig. 4.16
C, D). The archesporium is endothecial in origin. The cells of the endothecium divide and redivide
irregularly and form sporogenous cells. Large-sized sterile cells get differentiated at the base of the
capsular region of the sporogonium. Spiral thickenings develop on the walls of these sterile cells, and
this entire structure represents an elaterophore (Fig. 4.16). Some of the elaters remain attached also on
the elaterophore. The remaining sporogenous cells develop into fertile spore mother cells and sterile
elaters. Spiral thickenings develop on the walls of the elaters.
At the time of sporogenesis, each spore mother cell becomes a 4-lobed body. Its dipoid nucleus
divides meiotically to form four daughter nuclei, of which one each enters into each lobe (Fig. 4.16 H)
of the spore mother cell. Ultimately each uninucleate lobe develops into a haploid spore. In a majority
of the species, including Pellia epiphylla, the number of chromosomes in the gametophyte is nine
(n = 9).
Anticlinal divisions followed by periclinal divisions in the jacket initials form a jacket layer of two
or more cells thick. A sheath, called calyptra develops above the young sporogonium.
Seta remains short for quite some time. But soon it elongates rapidly. In some species (e.g. Pellia
epiphylla), the seta sometimes elongates so rapidly that from 1 mm it becomes as large as 80 mm within
a week’s time.
The mature sporogonium (Fig. 4.16 F) is made up of foot, seta and capsule. The foot consists of
parenchymatous cells. It is conical in shape with edges like a collar around the basal part of the seta.
The seta is made up of regular longitudinal rows of cells, which contain starch when young. The cells
of seta in the mature sporogonium are quite elongated and are devoid of starch grains. Capsule is a
globular or spherical structure surrounded by a jacket of two or more layer of cells. The outer wall layer
usually contains brown radial thickening bands while the cells of the inner wall layer contain many
semi-annular thickening bands in most of the species. An elaterophore is present in the lower end of the
base of the capsule. It consists of a bundle of stout fixed elaters, which are 20 to 100 in number. Elaters
are long, spindle-shaped bodies, each with 2 to 3 spiral thickened bands (Fig. 4.16 G). Thousands of
haploid spores are present in the capsule, along with elaters (Fig. 4.16F).
Dehiscence of capsule starts by an extraordinary elongation of the cells of the seta. They elongate
as much as 40 to 50 times. During this process of elongation, the starch of the seta cells converts into
sugar with the simultaneous rapid absorption of water. All this pushes the capsule through the calyptra.
The capsule dehisces by splitting in four valves (Fig. 4.16 I,J) along the lines of dehiscence. During this
process of dehiscence, the elaterophore comes out and gets exposed (Fig. 4.16J). The process of spore
dispersal is promoted by hygroscopic movement of elaters and elaterophore.
58 � Bryophyta
Fig. 4.16 A-E, Stages of the development of sporogonium in Pellia epiphylla; F, A mature sporogonium in
longitudinal sec on; G, An elater; H, A spore tetrad; I, A capsule showing dehiscence and exposed
elaters; J, A capsule showing lines of dehiscence; K, Germina on of mul cellular spore
Dispersed spores are multicellular, 4- to 9-celled, oval bodies (Fig. 4.16 K). Its basal portion extends
into a rhizoid. An apical cell gets differentiated in the apical region, and ultimately a new thallus of
Pellia is resulted.
A semi-diagrammatic depiction of the life cycle is shown in Fig. 4.17 (A-M).
Fig. 4.17 A-M. Semi-diagramma c representa on of the life cycle of Pellia
RICCARDIACEAE 4.11
Evans (1939) included only two genera under the family Riccardiaceae, These are Riccardia and
Cryptothallus. Some of the major characteristic features of Riccardiaceae are listed below.
1. Plant body is thalloid, and cells of the thallus contain prominent finely segmented oil bodies.
2. Sex organs are present on the dorsal side on short lateral branches.
3. The capsule is ovoid to cylindrical and remains surrounded by calyptra.
4. The calyptra is thick, fleshy, large and quite prominent.
5. The involucre is absent.
6. Elaterophore is present but it is distal in position. It is made up of long prosenchymatous cells,
marginal ones of which have a free tip.
7. Few, fixed elater-like cells also grow from the surface of elaterophore.
8. The capsule dehisces longitudinally into four valves, extending towards its base.
60 � Bryophyta
Some details of the life history of Riccardia are discussed below.
RICCARDIA 4.12
Riccardia is represented by about 300 species, distributed throughout the world, but mainly in tropics
and subtropical regions. R. multifida and R. pinguis are almost cosmopolitan in their distribution. Ten
species reported from India are Riccardia cardoti, R. decolyana, R. foreavana, R. indica, R. leveiri,
R. multifida, R. palmatiformis, R. pinguis, R. sikkimensis and R. villosa, as also mentioned by Parihar
(1987). It grows luxuriantly in a variety of habitats including wet grounds, moist rocks, moist sandy
grounds, decaying wood, ditches, marshy habitats as well as ditches and rocks near streams. R. multifida
grows extensively in the eastern Himalayan regions and hills of south India while R. pinguis grows
commonly in western Himalayas and hills of south India.
Originally named as Riccardius by SF Gray, the name Riccardia was given to this genus later in
1870 by Carrington. Some bryologists treat Aneura as a synonymn of Riccardia.
The plant body (Fig. 4.18A-D) or gametophytes are either completely thalloid or their terminal
branches are thalloid. The gametophytes are either completely prostrate (e.g. Riccardia indica; Fig.
4.18 A, B) or bear upright shoots developing from their prostrate, rhizome-like portion. A distinct
midrib is usually absent in prostrate species. In R. multifida and a few more species, the thalli are
pinnately branched. Some ill-defined organs of attachment arise from the lowermost parts of plants.
Incurved margins of some species form water sacs-like organs of water retention. Smooth-walled
rhizoids and mucilaginous hairs are present on the ventral surface of the thallus in species with
prostrate thalli. The prostrate thallus may be thick, broad and slightly branched (e.g. R. pinguis),
or regularly branched or irregularly branched or even pinnately branched (e.g. R. multifida).
Fig. 4.18 A-G Thalli of some species of Riccardia. A-B, R. indica; C-D, R. sikkimensis;
E, R. prehensilis; F-G, R. pinguis
Anatomically, the thallus lacks any differentiation of tissues and consists of 5 to 15 layers of cellls
with cells of superficial layers being comparatively smaller. Thallus is several layers thick in the middle
and becomes narrower to even one-cell layer towards the margins. Cells, when young and exposed
to sunlight, contain chloroplasts. Some oil bodies are also present in epidermal cells. The number of
oil bodies in each epidermal cell varies from 3 to 40 in different species. In a few species only (e.g.
R. prehensilis), there is some internal differentiation of tissues showing a region of even some thick-
walled cells (Fig. 4.19C).
Apical growth takes place by a wedge-shaped apical cell with two cutting faces. It alternately cuts
off segments on right and left sides. However, according to showalter (1923), two-faced apical cell
“cuts off segments not to the right and left but below and above.
Vegetative reproduction takes place (i) community by progressive death and decay of the older parts
of thallus, separating the branches, which develop into new thalli; (ii) by producing stolons or thick
cylindrical ventral shoots, which on separation produce new thalli, as in Riccardia levieri (Pande and
Srivastava, 1958); and (iii) by producing gemmae at the apex of branches in several species, including R.
levieri, R. multifida and R. palmata. Gemmae in Riccardia are small, round, oval or oblong bodies only
of few cells. On being detached and germination, each gemma produces a new thallus of Riccardia.
Sexual reproduction in Riccardia is oogamous like other bryophytes, and takes place by antheridia
and oogonia. Species may be monoecious (e.g. R. decolyana and R. multifida) or dioecious (e.g. R.
indica and R. palmata). Sex organs develop on specialized short lateral branches, which develop from
the margins of the thallus.
Antheridia develop on dorsal surface of thallus on lateral branches (Fig. 4.18 B,C) called male or
antheridial branches. They remain sunk in the antheridial chambers in the tissues of male branches
(Fig. 4.20 A, B). The development of antheridium follows the same pattern as described for Porella
and other Jungermanniales under Article 4.5.9(1) (Fig. 4.3).
Fig. 4.19 A-C. Transverse sec ons of the thallus of Riccardia mul fida (A),
R. indica (B), and part of axis of R. prehensilis (C)
62 � Bryophyta
Fig. 4.20 A-B, Antheridia sunken in antheridial branch (A) and a single antheridium (B) of Riccardia
The mature antheridium is a globular body with a very short stalk of only 2 or 3 cells in length. It
is covered by a single-layered jacket (Fig. 4.20 B). Androcytes present inside the jacket develop into
antherozoids. Each antherozoid bears two short flagella.
Archegonia develop on the dorsal surface of the archegonial branches (Fig. 4.18E), which
are comparatively shorter than the antheridial branches. Each archegonial branch possesses 4-20
archegonia arranged usually in two alternate rows. The development of archegonium resembles other
Jungermanniales as discussed for Porella under Article 4.5.10(1) (Fig. 4.5 A-I).
The mature archegonium (Fig. 4.21 A-B) shows only a little differentiation into neck and venter.
Usually, the venter as well as the lower part of neck are 2 or 3 cells in thickness. The neck consists
of five vertical rows of cells, with its axial row made up of 3 to 6 neck canal cells, a ventral canal
cell and an egg (Fig. 4.21C). Involucral scales surround and protect the lower parts of the developing
archegonia. The fertilization process also resembles that of Porella and other Jungermanniales, as
discussed earlier under Article 4.5.11. Fusion of male and female gametes results in the formation of
diploid zygotes.
Fig. 4.21. A, An enlarged female branch of Riccardia diversifolia; B, Sec on through

an archegonial branch of R. diversifolia; C, A mature archegonium of R. sinuata
Development of sporogonium starts by a first transverse division (Fig. 4.22A) of the zygote resulting
into an epibasal cell and a hypobasal cell. The entire sporogonium develops from the epibasal cell. The
lower hypobasal cell simply elongates and develops into a haustorium (Fig. 4.22B), which pierces into
the gametophytic tissue. The epibasal cell divides transversely to form two cells, both of which contain
prominent nuclei, dense cytoplasm and many chloroplasts. A transverse division in the uppermost
cell makes the young sporogonium a four-celled structure, of which the lowermost is the haustorium.
Of the three cells formed by the epibasal cell, the lowermost develops into foot, middle one into seta,
and the uppermost into capsule of the sporogonium. Intersecting vertical divisions in these three cells
result into the formation of three superimposed tiers, each made up of four cells (Fig. 4.22C). The tier
immediately next to haustorium develops into foot. Its cells divide by transverse as well as vertical
divisions in an irregular fashion and form a compact mass of foot. Vertical and transverse divisions in
the cells of middle tier from seta. The uppermost tier of four cells develops into capsule. A transverse
division in this uppermost tier divides this tier into two tiers of four cells each. Now periclinal division
in each cell of these two tiers results into the formation of an outer or peripheral amphithecium and inner
or axial endothecium (Fig. 4.22D). The amphithecium develops into the jacket of the capsule. Its cells
divide first anticlinally and then also periclinally to form a two-layered thick jacket (Fig. 4.22 E, F).
The archesporium is endothecial in origin (Fig. 4.22D). Endothecial cells divide irregularly. The cells
of apical central region originated from endothecium are larger and develop into elaterophore (Fig.
4.22F). From these cells radiate the sporogenous tissue, which later on differentiate into spore mother
cells and elaters (Fig. 4.22F). Prior to nuclear division, the spore mother cells attain a four-lobed
structure. The diploid nucleus of each spore mother cell divides by two successive divisions to form a
tetrad of haploid spores. A massive sheath of gametophytic tissue keeps enclosing the young capsule.
At maturity, this sheath breaks and survives at the basal part of the seta in the form of a collar.
The mature sporogonium of Riccardia (Fig. 4.22 F) is made up of foot, seta and capsule. The foot
is ill-defined and present in the form of a club-like swelling in the lowermost part of the sporogonium.
Seta consists of several rows of cells, having a diameter of at least six cells. The capsule is a cylindrical
to ovoid body (Fig. 4.22F) surrounded by a bilayered jacket. Bands of thickenings are present in the
walls of the cells of jacket of most species of Riccardia. A well-developed apical elaterophore hangs
downwards into the cavity of the capsule up to as much as one-third of the distance from the apex to
the base. It forms a central cylinder of long parenchymatous cells. The marginal cells of elaterophore
have free tips. Fixed elaters or elater-like cells grow from the surface of the elaterophore. These project
from the mass of free elaters and spores. A prominent spiral band of thickening is present in each free
elater (Fig. 4.23 A). Ends of the free elaters are pointed.
At the time of the dehiscence of a capsule, cells of the seta start elongating. Seta thus elongates
from 2 to 30 mm. Along the line of dehiscence, the capsule dehisces into four valves, which separate
out from apex to base. The elaterophore also splits into four parts during this process. The mechanical
jerk results into direct and abrupt dispersal of spores. Hygroscopic movement of elaters also helps in
spore dispersal.
The spores germinate into gametophytes (Fig. 4.23 B-D). They are small, 12 to 25 m in diameter,
and each remains surrounded by two wall layers, namely exospore and endospore. A nucleus and
chloroplasts are also present in each spore. At the time of germination, the spore increases in size,
divides transversely and forms two equal or unequal cells. An oblique division in the upper cell
differentiates a wedge-shaped apical cell, the further activity of which gives rise to a new gametophytic
thallus (Fig. 4.23 D).
64 � Bryophyta
Fig. 4.22 A-F. Showing development of sporogonium in Riccardia pinguis

Fig. 4.23 A-D, Riccardia levieri, showing an elater (A) and spores (B) and development of young gametophyte
FOSSOMBRONIACEAE 4.13
Fossombroniaceae is one of the seven familes of Metzgerineae. It includes four genera, viz.
Fossombronia, Simodon, Petalophyllum and Sewardiella. Majority of the species of Fossombroniaceae,
including of Fossombronia, have gametophytes differentiated into stem and lateral leaves. Growth of
the gametophyte is initiated by a single apical cell.
Only Fossombronia is discussed here in some details.
FOSSOMBRONIA 4.14
Fossombronia is represented by about 50 species showing worldwide distribution. Four of its species
reported from India are F. indica, F. himalayensis, F. cristula and F. foreavii. In India, Fossombronia
has been reported from south India, Himalayas, Pachmarhi (MP) and Kodaikanal.
The gametophytic plant body is foliose. The thallus is almost completely prostrate with a sparse
or profuse branching (Fig. 4.24 A–C). Each branch is made up of a well-developed stem bearing a
single row of leaves along both the lateral margins. The arrangement of leaves becomes less organised
in some species because of the greatly convoluted margins of their leaves (Fig. 4.24 A). The leaves
are 2- to 3-cells thick at the base while their upper portions are only one cell in thickness. The well-
developed massive stem shows no indication of any internal differentiation of tissue. Only smooth-
walled rhizoids are present on the ventral surface of stem. The apex of the branch contains many
multicellular mucilaginous hairs. Their function is to protect the apical region. Older parts of the stem
do not bear any mucilaginous hairs.
Growth takes place by an apical cell with two cutting faces. It cuts segments alternately on the
right and left sides. The apical cell is semicircular in outline in vertical longitudinal section. It appears
narrowly trianglular in HLS. A segment of an apical cell is first cut in a horizontal plane. The ventral
daughter cell, formed by this division, gives rise to the ventral portion of the stem while the dorsal
daughter cell gives rise to the dorsal portion of the stem and also the leaf. Differing from other members
of Metzgerineae, the branching in Fossombronia is not initiated by a vertical division of the apical cell
66 � Bryophyta
into two equal-sized cells. In this genus, a second apical cell is differentiated in a very young segment
derived from the apical cell. The overall growth from the original apical cell and this second apical cell
ultimately gives rise to a structure which appears to be a true dichotomy. In reality, however, it is a false
dichotomy and not a true dichotomy.
The sex organs develop in acropetal succession on the dorsal surface of the midrib and occur either
irregularly scattered or in groups. A majority of the species are monoecious. According to Pande et al.
(1954), Fossombronia himalayensis is monoecious but protandrous. Rarely, some species are dioecious.
Each sex organ generally originates from a single, superficial cell, quite close and just behind the apical
cell.
Development of antheridia follows the same pattern as that of other Jungermanniales and explained
already for Porella under Article 4.5.9(1). Stages of the development of antheridia in Fossombronia
angulosa are depicted in Fig. 4.25A-G. Mature antheridia burst suddenly and release the antherozoids.
The antherozoids are biflagellate, uninucleate structures. Two flagella of each antherozoid remain
inserted at different points near the anterior end. Mature antheridium (Fig. 4.25 G) is a stalked structure
and contains a globular body surrounded by a jacket layer. The haploid chromosome number in the
dividing antheridial cells of F. himalayensis is nine (Pande et al., 1954).
Fig. 4.24 Gametophytes of Fossombronia. A, Male gametophyte of F. longiseta; B, Female

gametophyte of F. longiseta; C, Female gametophyte of F. intes nealis
Fig. 4.25 A-G, Stages of the development of antheridia in Fossombronia angulosa; H-R, Stages of the
development of archegonia in F. angulosa. Stages in J, Q and R are in transverse sec ons
Development of archegonium (Fig. 4.25 H-R) is also in the manner typical for the Jungermanniales,
as described for Porella under Article 4.5.10(1). A mature archegonium (Fig. 4.25O) consists of 6-8
neck canal cells surrounded by five vertical rows of neck cells. The venter of the archegonium contains
a two-cells thick jacket layer. The venter is only slightly broader than the neck of the archegonium.
At the time of fertilization, entrance of antherozoids into archegonia is undoubtedly in response
to chemotactic stimuli. Antherozoids entering the venter of an archegonium soon penetrate the egg.
Fusion of male and female nuclei takes place 24 to 48 hours after the entrance of the antherozoid, and
it results into the formation of a diploid zygote.
68 � Bryophyta
According to Showalter (1927), the division of the zygote “takes place six to nine days after gametic
union. The zygote first divides transversely into a larger epibasal cell and a smaller hypobasal cell
(Fig. 4.26 A). The hypobasal cell develops into the foot of the sporophyte. The epibasal cell divides
transversely into an upper daughter cell, which develops into the capsule, and a lower daughter cell,
which develops into a seta of the sporophyte. (Fig. 4.26 B-E). Periclinal divisions in the capsular
region result in the formation of an outer jacket layer or amphithecium and inner endothecial cells. The
archesoporium is endothecial in origin.
Fig. 4.26 A–E, Stages of the development of embryo in Fossombronia pusilla
The jacket or amphithecium soon becomes a two-celled thick layer. Elaters and sporocytes do not
differentiate until the last cell generation of the sporogenous tissue of the endothecium. The elaters of
Fossombronia are unique in that the thickenings on their walls are laid down in 5 to 9 rings instead of
the usual 2 or 3 longitudinal thickenings of the other genera. Elaterophore-like structure, when present,
is ill-developed.
A calyptra and a cup-like involucre enclose the young sporophyte. Calyptra is actually the part
of the old archegonium, while the involucre is produced by the upgrowth of the gametophytic tissue
adjacent to the basal part of the archegonium. Until the spores in the capsule attain maturity, the seta
is only about 1 mm in length. Soon, it elongates rapidly and pushes the capsule through the calyptra
(Fig. 4.24B). After the elongation of seta, the capsule dehisces after about 18 to 36 hours. Capsules in
different species either dehisce irregularly or into four valves.
The dehisced spores start germinating in the form of a filamentous protonema of 2 to 12 cells.
Rhizoids start developing in the cells nearest the old spore wall. Some distal cells of the protonema start
dividing irregularly to form a globose mass of cells. This is actually the termination of the protonemal
phase. A cell of this globule mass of cells soon starts functioning as an apical cell with two cutting
faces. A thalloid structure soon develops due to the activity of this apical call. There is, however, still no
differentiation of stem and leaves. At a later stage, derivatives of the apical cell start differentiating into
a leaf and a portion of the stem. Within few days, a foliose gametophyte of Fossombronia develops.
1. Name the largest order of Hepa copsida.

2. Write any seven general characteris cs of Jungermanniales.
3. Are the rhizoids in Jungermanniales smooth-walled or tuberculate or of both types?
4. Are the scales present or absent in Jungermanniales?
5. Verdoorn (1932) divided Jungermanniales into two ar ficial groups. These are (a)
Jungermanniales Acrogynae, and (b) _____ _____.
6. Evans (1939) divided Jungermanniales into three sub-orders. These are (a) Haplomitrineae,
(b) Metzgerineae, and (c) _____.
7. Parihar (1987) divided all Jungermanniales into two sub-orders. These are (a) Jungermannineae,
and (b) _____.
8. Write at least five characteris c features of the sub-order Jugermannineae (Jungermanniales
Acrogynae).
9. Porellaceae and Frullaniaceae belong to the sub-order _____ of the order Jungermanniales.
10. Porellaceae is also known by the name _____________.
11. Write at least four characteris c features of Porellaceae.
12. What do you mean by incubuous arrangement of leaves?
13. Draw well-labelled diagrams of external morphology of the gametophyte of Porella.
14. Explain the following terms in reference to the leaf of Porella:
(a) Amphigastria
(b) An cal lobe
(c) Pos cal lobe
(d) Incubous
15. Describe life-history details of Porella in about 500 words.
16. Give an illustrated account of development of antheridium in Porella.
17. Explain in brief the generalised process of archegonium development in Porella and other
Jungermanniales.
18. Describe development of sporogonium and structure of mature sporogonium in Porella.
19. Three genera included under Frullaniaceae are Frullania, Neoha oria and ____________.
20. Write at least four characteris c features of Frullaniaceae.
21. Write a detailed scien fic note on the life-history of Frullania in about 500 words.
22. The common name “mul form thallose hepa cs” is usually given to the members of
Metzgerineae. Why?
23. Describe distribu on and habitat of Pellia in about 100 words.
24. Give an illustrated account of life-history of Pellia in about 1000 words.
25. Describe the development of sporogonium and structure of mature sporogonium in Pellia.
26. What is elaterophore?
27. In Pellia, the archesporium is _______ in origin.
28. Draw a semi-diagramma c sketch of the life cycle of Pellia.
29. Two genera included under the family Riccardiaceae are Riccardia and __________.
30. Write a detailed scien fic note on the life history of Riccardia giving suitable diagrams.
31. Give a brief illustrated account of the life history of Fossombronia.
5
Hepaticopsida
(Sphaerocarpales)
SPHAEROCARPALES AND THEIR GENERAL CHARACTERISTICS 5.1
Sphaerocarpales is a small order of Hepaticopsida, comprising only three genera, viz. Sphaerocarpos,
Geothallus and Riella. The former two (i.e. Sphaerocarpos and Geothallus) belong to the family
Sphaerocarpaceae while the third one (i.e. Riella) is the sole member of the family Riellaceae.
Members are commonly called “bottle-hepatics” because of the presence of a flask-shaped
or globose envelope (involucre) around each of the sex organs. Sphaerocarpaceae members have
bilaterally symmetrical and thallose gametophytes with no internal differentiation of tissues. On the
other hand, Riellaceae (Riella) members possess asymmetrical gametophytes. A well-organised seta is
absent in the sporophyte of the members of Sphaerocarpales.
Some details of the life-history of Sphaerocarpos of the family Sphaerocarpaceae and Riella of
Riellaceae are given here in this account.
SPHAEROCARPOS 5.2
Sphaerocarpos is represented by seven species, and occurs in both northern as well as southern
hemispheres. It occurs commonly in the Gulf and Pacific coast states of the United States on fine-
textured soil. It prefers a climate where the summer is dry and winter is moderately wet and wild.
The plant body is gametophytic and gametophytes are relatively small, orbiculate to cuneate and
quite simple or slightly dichotomously branched. Gametophyte is a bilaterally symmetrical thallus.
Male and female gametophytic plants (Fig. 5.1 A-B) usually occur together in clumps. A several cells
thick, broad midrib is present in each gametophytic thallus. On margins, the wings of the thallus are
only one cell in thickness. Wings of the thallus are either entire or incised into leaf-like lobes. Sex
organs, each surrounded by an ovoid or flask-shaped involucre, are present on the dorsal surface of a
midrib. They remain densely crowded (Fig. 5.1). On the ventral surface of the thallus are present many
multicellular glandular hairs and smooth-walled rhizoids. Tuberculate rhizoids and scales are absent.
Anatomically, the thallus lacks any internal differentiation of tissues. Chloroplasts remain filled in
almost all vegetative cells of the thallus, except rhizoids.
Hepaticopsida (Sphaerocarpales) � 71
Fig. 5.1 Female (A) and male (B) gametophytes of Sphaerocarpos californicus; C-I, Stages of the
development of antheridia in S. cristatus; J-M, Stages of the metamorphosis of androcytes into
antherozoids in S. donnellii
Growth takes place by a row of wedge-shaped apical cells, which remain laterally joined to one
another. Each apical cell cuts off segments alternately from its dorsal as well as ventral faces. These
segments contribute to the midrib of the thallus. The segments, which are cut off from the apical cell
at each end of the row, ultimately form wings of the gametophytic thallus. Vertical divisions in the
median part of the row of apical cells ultimately contribute to the dichotomous branching of the thallus.
Repeated dichotomous branching is common in species like Sphaerocarpos donnellii.
Vegetative reproduction takes place by progressive death and decay of the posterior portion of the
thallus. When this death and decay process reaches up to the place of dichotomy, the two branches may
survive and may continue to develop into two independent plants of Sphaerocarpos. Sometimes, plants
multiply vegetatively also by the formation of proliferous outgrowth, either from the midrib or from
the lateral wings. Such outgrowths may also develop from the involucres in some species, according
to Rickett (1920).
72 � Bryophyta
Sexual reproduction is oogamous. All the investigated species of Sphaerocarpos are heterothallic.
Sex differentiation is attributed to sex chromosomes. Male plants (Fig. 5.1 B) differ from female
plants by their (i) smaller-sized gametophytes, (ii) flask-shaped involucres, and (iii) purple-tinged
gametophytes. Male plants attain a diameter of about 2 mm while the female plants reach up to 1 cm
in diameter. In female plants, the archegonia are surrounded by subspherical or cylindrical involucres
(Fig. 5.1A).
Development of antheridium (Fig. 5.1 C-I) starts with a superficial dorsal cell (Fig. 5.1C), which
enlarges and becomes capitate. It is called antheridial initial. It soon divides into a basal cell and an
outer cell. The outer cell projects above the thallus (Fig. 5.1 D). The basal cell develops into antheridial
stalk. The outer cell develops into the remaining major part of the antheridium. The outer cell divides
by successive transverse divisions to form a vertical file of three cells (Fig. 5.1E), of which two upper
cells are the primary antheridial cells and develop further into antheridium proper. The lowermost
third cell forms the primary stalk cell. Each of the two primary antheridial cells divide by two
successive vertical divisions at right angles to each other and ultimately forms four cells (Fig. 5.1 F).
A periclinal division in both tiers of four cells results into the formation of outer four jacket initials
and inner four primary androgonial cells (Fig. 5.1.G). Primary androgonial cells now divide by many
successive divisions at right angles to one another to form many cubical androgonial cells. Each cell
of the last cell generation of the androgonial cells divides diagonally into two androcytes. The jacket
initials divide anticlinally to form a well-organised jacket around the antheridium (Fig. 5.1I). Each
androcyte metamorphoses into a uninucleate, biflagellate antherozoid (Fig. 5.1 J-M). Free swimming
antherozoids are spindle-shaped, curved or variously coiled structures.
Development of archegonium starts with the arrangement of a dorsal cell called archegonial
initial (Fig. 5.2A). It first divides by a transverse division into a basal cell and an outer cell (Fig.
5.2B). The lowermost part of the archegonium is formed by the basal cell, while its remaining portion
is formed by the outer cell. The outer cell first divides by an asymmetrical vertical division to cut
off a peripherial initial. Two more unequal vertical divisions form two more peripheral initials. A
primary axial cell surrounded by these three peripheral initials now develops (Fig. 5.2 C). All the three
peripheral initials now divide vertically to form six jacket initials surrounding the primary axial cell.
A transverse division in each jacket initial soon results in the production of six neck initials present
above a tier of six venter initials (Fig. 5.2D). An archegonial neck of several cells in length and six
cells in perimeter now develops due to transverse division of neck initials and their daughter cells. In
Sphaerocarpos, the jacket of a mature venter of archegonium consists of 10 or more cells in perimeter.
Simultaneously, the primary axial cell divides transversely into a primary cover cell and a central cell
(Fig. 5.2D). The primary cover cell divides by two successive vertical divisions to form four cover
cells. Side by side, the central cell divides transversely into a smaller upper canal cell and a lower
canal cell (Fig. 5.2 E). The upper canal cell divides by two successive transverse divisions resulting
into the formation of four neck canal cells within the archeogonial neck. The lower canal cell now
divides asymmetrically to form a small venter canal cell and a large egg. Soon, it is observed that the
venter canal cell and neck canal cells of the mature archegonium disintegrate to form a mucilaginous
mass, and its cover cells are set apart, resulting in the formation of an open passage for permitting and
helping the entry of the antherozoids to reach up to the egg. Entrance of antherozoids “into the neck is
undoubtedly a chemotactic response” (Smith, 1955).
An involucre starts developing in the initial stages of the archegonial development (Fig. 5.2 C-G).
The involucre is more prominent around the antheridium (Fig. 5.1 E-I) than that of the archegonium
(Fig. 5.2 G).
Fig. 5.2 A-M Sphaerocarpos cristatus. Development of archegonia (A–G); Development of

sporophyte (H-L); M, Mature sporophyte
Fertilization takes place due to the fusion of antherozoids and the egg. Fusing male and female
nuclei have organised chromosomes, and fusion of male and female nuclei begins 60 to 70 hours after
entrance of an antherozoid. The zygote first divides transversely to form an upper epibasal and a lower
74 � Bryophyta
hypobasal cell, and both these cells again divide transversely to form a 4-celled filamentous embryo
(Fig. 5.2H). Each of the cells of this young embryo divide by two successive vertical divisions to
develop into a tier of four cells. Two tiers of the four cells in the epibasal half of the embryo develop
ultimately into a capsule of the sporophyte while the derivatives of the remaining two tiers develop
into the remaining parts of the sporophyte. Periclinal division (Fig. 5.2I, J) in the two upper tiers of
four cells results into the formation of outer amphithecium and inner or central endothecium. A jacket
layer of capsule develops due to anticlinal division in the amphithecium (Fig. 5.2 J-M). Archesporium
is endothecial in origin in Sphaerocarpos. The endothecium develops into sporogenous tissue of the
capsule. Some sporocytes or spore mother cells of the sporogenous tissue divide meiotically into four
spores. Other cells of the sporogenous tissue remain sterile and mature into sterile nurse cells. The
function of nurse cells is to supply food to the developing spores. Numerous chloroplasts are present in
the nurse cells as well as in the cells of the jacket layer of the capsule. Dispersal of spores takes place
after the death and decay of the jacket.
A bulbous foot develops simultaneously by the divisions in the basal part of the hypobasal half of
the embryo (Fig. 5.2 L,M). The upper part of the hypobasal half of the embryo develops into a small
seta. Seta in Sphaerocarpos, however, does not elongate much. It does not also pushes much to the
well-developed capsule of the sporophyte.
Out of the four spores formed by a spore mother cell, two develop into female gametophytes, and two
into male gametophytes. A spore first geminates into a long germ tube, which first divides transversely
into a small terminal cell and a large basal cell. The terminal cell remains filled with dense protoplasm
and divides by transverse and vertical divisions to form a long ribbon-shaped structure of 3 to 4 cells
in length and two cells in breadth. The basal cell does not divide any further. Horizontal divisions in
the multicellular ribbon give rise to the formation of a germinal disc, which soon grows and forms an
asymmetrical structure. Apical cells develop at one side of this disc. Their further activity results into
the development of an adult thallus of Sphaerocarpos.
RIELLA 5.3
Riella is the only genus of the family Riellaceae of Sphaerocarpales. It differs from Sphaerocarpaceae
in possessing asymmetrical plant body of its gametophytes. About 20 species of Riella have been
reported from the world, of which R. affinis and R. americanum grow commonly in the United States.
R. affinis is the only species reported from India.
Riella is an aquatic bryophyte, growing on muddy soil, which remains submerged in calcium-rich
waters of shallow pools. Riella plants grow entirely submerged, usually several feet below the water
surface.
The gametophytic plant body (Fig. 5.3 A) of Riella is erect and consists of a thick stem-like axis
bearing a well-developed, plate-like wing. The wingh is straight to spirally twisted and only one cell in
thickness. The cells of the wing are thin-walled and chlorophyllous. One-cell-thick scales (Fig. 5.3 B)
develop along the median part of the axis. These are called ventral scales. Some scales also develop
at the juncture of the axis and wing. These are called lateral scales. Simple spherical oil bodies are
present in some cells of the scales. The lower part of the thallus contains some unicellular and smooth-
walled rhizoids.
Growth takes place by a single apical cell with two cutting faces. Sometimes the single apical cell
is replaced by a row of apical cells.
Fig. 5.3 Riella. A, Gametophyte with sporophytes of R. americana; B, Thallus with scales and gemmae; C, A
part of the thallus with antheridia on the margin; D, A part of thallus with archegonia; E-F, Young
antheridia of R. affinis; G-H, Young and mature archegonia of R. affinis; I, A sporophyte of R. affinis
Vegetative reproduction takes place by gemmae (Fig. 5.3 B), which develop between the rows
of lateral and ventral scales. A mature gemma is a one-cell thick, spherical or oval structure. It is
transversely constricted to two unequal-sized lobes. Studhalter and Cox (1940, 1941) called gemma of
Riella as a gemmaling. On being detached, a gemmaling becomes meristematic in a place between two
lobes and develops into an adult gametophytic phase.
76 � Bryophyta
Riella may be homothallic or heterothallic. Antheridia always develop in clusters in notches along
the margin of the wing of gametophytic thallus. Archegonia develop singly at the juncture of the axis
and wing.
Thompson (1943) studied the development of sex organs in Riella (Fig. 5.3 E–H) and observed that
it is quite similar to that of Sphaerocarpos, described earlier under Article 5.2 (Fig. 5.1 C-I and 5.2
A-G). Minor deviations in the development of archegonial involucres, however, exist. Here, in Riella,
the archegonial involucre originates from the stalk of the archegonium.
Thompson (1942, 1943) also studied the development of sporophyte in Riella and reported that it is
also quite similar to that of Sphaerocarpos. Sporocytes and quadrinucleate nurse cells also develop by
the sporogenous tissue of the sporophyte of Riella (Fig. 5.3I).
The haploid spore germinates in the form of a long germ tube, at the tip of which differentiate two
apical cells. A series of short broad cells result due to the transverse divisions in these apical cells. These
cells now divide longitudinally to form a small but flat thallus. Now the active cell division process in
the juvenile phase is restricted only up to the margins, and soon a single apical cell is differentiated. Its
activity results in the production of an adult phase of gametophyte bearing axis and wing.
1. Sphaerocarpales include only three genera, namely Sphaerocarpos, Riella and _____.
2. The members of Sphaerocarpales are commonly called _____.
3. Why is the name “bo le-hepa cs” given to members of Sphaerocarpales?
4. Give only one characteris c feature on the basis of which one can differen ate between
Sphaerocarpaceae and Riellaceae.
5. Describe in brief some life-history details of Sphaerocarpos in about 500 words.
6. Draw neat and well-labelled diagrams of the following:
(a) Male gametophyte of Sphaerocarpos
(b) Female gametophyte of Sphaerocarpos
(c) A mature sporophyte of Sphaerocarpos
(d) A mature sporophyte of Riella
7. Is Riella an aqua c bryophyte or a terrestrial bryophyte?
8. In Sphaerocarpos, the archesporium is _____ in origin.
9. In Sphaerocarpos, is seta the very small or very long?
10. Write a short scien fic note on Riella in about 200 words.
11. Draw a labelled diagram depic ng external features of the gametophyte of Riella bearing
sporophytes.
6
Hepaticopsida
(Monocleales)
WHAT ARE MONOCLEALES? 6.1
As mentioned earlier under Article 3.4, Monocleaceae is a unigeneric family which shows characters of
both Marchantiales and Jungermanniales, and has been placed by some bryologists under Marchantiales
and by others amongst Jungermanniales. Due to Calobryum-type of archegonial development,
Monocleales show closeness to the order Calobryales. It was Schuster (1963), who on the basis of
his detailed studies of antipodal Hepaticae, suggested it to be placed in a separate order Monocleales,
including a single family Monocleaceae and a single genus Monoclea. Schuster (1963) and later on
Sandra Holmes (1986) treated Monocleales as an order of Hepaticopsida.
GENERAL CHARACTERISTICS OF MONOCLEALES 6.2

1. Monocleales is a monotypic order represented by a few species of its only genus Monoclea.
2. Plant body is thalloid and remarkably large, reaching up to 20 cm or more in length and 5 cm
or more in breadth, and that is why these are commonly called “giant-thallose-liverworts”.
3. Monocleales resemble members of the order Metzgeriales in possessing several characters
like (i) lack of air chambers in the gametophyte, (ii) lack of ventral scales, (iii) long seta,
(iv) elongate elaters, (v) elaborate calyptra, and (iii) an archegonial pouch.
4. Monocleales also resemble members of the order Marchantiales in possessing several similar
characters like (i) specialised antheridial pads, (ii) embedded antheridia, (iii) unlobed spore
mother cells, (iv) presence of oil bodies in isolated cells, (v) presence of two types of rhizoids,
(vi) structure of neck of archegonia, and (vii) structure and dehiscence of the jacket of the
capsule.
Characters of Monocleales mentioned above under (3) and (4) suggest these members have
an independent position as a separate evolutionary line, and hence an independent order of
Hepaticopsida.
5. Monocleales differ from Marchantiales in possessing the unique hood-like sheath posterior to
the female receptacle.
6. Monocleales also differ from Marchantiales in possessing the elongate capsule.
78 � Bryophyta
MONOCLEA 6.3
Monoclea is the sole representative of the only family (Monocleaceae) of order Monocleales. Two of
its species reported so far are M. gottschei and M. forsteri. M. gottschei has been reported from tropical
America while M. forsteri from tropical America as well as New Zealand.
Smith (1995) stated that “twice to thrice dichotomously branched gametophyte of Monoclea
forsteri (Fig. 6.1A) is the largest known thalloid gametophyte among Bryophyta”, and that is why
these members are commonly called “giant-thallose-liverworts”. Thallus lacks a clear midrib. The
plant body is only 1 or 2 cells thick on the margins while it reaches up to 10 cells or more in thickness
in the middle. The dorsal surface of the thallus is smooth. Rhizoids are confined only on the ventral
surface of the thallus. They are smooth-walled as well as some also contain irregular thickenings and
thus resemble tuberculate rhizoids. Scales are absent. Growing apex of the thallus is protected by some
unicellular mucilaginous hairs.
Internally, the thallus is parenchymatous and remains covered by a layer of dorsal epidermis
and ventral or lower epidermis. Cells of the dorsal or upper epidermis and few cells immediately
beneath it are filled with chloroplasts. Few cells near the upper epidermis also contain some crystals
of calcium oxalate. Walls of some of the cells also have conspicuous pores, which probably provide
internal connection for transport of water. Air chambers, characteristic of Marchantiales, are absent in
Monoclea. Some parenchymatous cells remain filled with oil bodies. Lower or ventral epidermis bear
both smooth-walled and tuberculate rhizoids, and no scales.
Reproduction in Monoclea resembles Riccia, specially the ontogeny of antheridium as well as
archegonium. Monoclea is heterothallic. The antheridia of each receptacle are produced in acropetal
succession. They remain enclosed in semicircular to elongate, cushionlike pads called receptacle,
which is present behind the growing point. Continuous growth and succession of male receptacles
is observed because the apical initial is not utilised in their formation. Each antheridium develops in
an antheridial chamber (Fig. 6.1B), which opens to the exterior by a long narrow canal. The mature
antheridium is an ovoid shortly-stalked structure, somewhat pointed towards the apex. Each antheridial
chamber opens on the dorsal surface by a pore.
The archegonia develop within a pouch-like, tubular structure called involucre (Fig. 6.1 C). The
involucre develops in the form of a hood-like outgrowth covering the archegonia. It may attain a
length of as much as 1.5 cm. Archegonia remain situated behind growing points of thallus. Monoclea
resembles Riccia in ontogeny of its archegonium. A mature archegonium contains a swollen venter
and a long neck of six vertical rows of jacket cells, and thus resembles Riccia as well as Marchantia.
The neck contains as many as 10 neck canal cells. Since production of archegonia does not check apical
growth in Monoclea, it is anacrogynous.
Two, three or even more sporophytes may be observed emerging from an involucral cavity (Fig.
6.1A). A young sporophyte also remains surrounded by a long calyptra (Fig. 6.1C). A mature sporophyte
possesses a long and massive seta reaching up to a length of as much as 4 cm. At the top of the seta
is present a dark-brown cylindrical capsule, which is broader than seta and attains a diameter of as
much a 1.5 mm. Great seta length and elongate capsule in the mature sporophyte are the characters of
Monoclea which bring it more close to Jungermanniales than that of Marchantiales.
Diploid zygotic nucleus divides first by many free nuclear divisions, and walls start appearing
slightly at a later stage. Foot, seta and capsule become quite clear during the early embryogeny. A
Hepaticopsida (Monocleales) � 79
single-layered amphithecium soon gets differentiated by periclinal division in the region of capsule.
The sporogenous tissue is endothecial in origin. Elaters, having 2 or 3 helical thickenings, soon start
differentiating in the sporogenous tissue. The fertile sporogenous cells form 8 isodiametric sporocytes,
each of which divides reductionally to form 4 haploid spores.
Fig. 6.1 A, A gametophyte of Monoclea forsteri bearing many sporophytes; B, Longitudinal ver cal sec on
through a male gametophyte of M. go scheri; C, Longitudinal ver cal sec on through a female
gametophyte of M. go escheri bearing a young sporophyte surrounded by calyptra and involucre;
D, A part of the capsule wall with forked transverse thickenings
80 � Bryophyta
The wall of the capsule is single-layered and contains forked transverse thickenings (Fig. 6.1), a
unique feature of Monoclea. Dehiscence of capsule takes place by a single longitudinal slit, through
which elaters uncoil abruptly and help in dispersal of spores. It never dehisces into four valves. Each
spore is a very small structure. It starts germinating almost immediately after dispersal. A multicellular
mass of cells is produced, from which the rhizoids start developing soon. An apical cell soon develops,
the activity of which results in the formation of a young gametophyte. Sporocytes in Monoclea also get
partitioned into four parts by furrowing, and this type of cytokinesis is seen commonly in members of
Jungermanniales. In Marchantiales, this type of cytokinesis is rarely seen.
1. What are Monocleales?

2. Monocleales shows characters of both Jungermanniales and _____.
3. Who was the first to suggest that Monocleaceae should be placed under an independent
order Monocleales?
4. How many genera do Monocleales include?
5. Write any four resemblances of Monocleales with Marchan ales.
6. What is the common name of Monocleales?
7. Write a detailed note on the life history of Monoclea in about 500 words.
8. In Monoclea, are scales present or absent?
9. Write a note on involucre of Monoclea in about 100 words.
10. Describe sporophyte of Monoclea in about 200 words.
7
Hepaticopsida
(Marchantiales)
LIMITS OF MARCHANTIALES? 7.1
Marchantiales is the most prominent order of the class Hepaticae or Hepaticopsida (Rothmaler,
1951). As mentioned earlier under Article 3.4, “recent studies of molecular biology, genosystematics,
phylogeny, diversification and classification of bryophytes (Newton et al., 2000; Norris, 2003; Shaw
and Renzaglia, 2004; Zander, 2006; and Triotsky et al., 2007) treat all bryophytes under an independent
subkingdom (Bryobionta) and divided it further into three phyla, of which one is Marchantiophyta
(= Liverworts or Hepaticopsida or Hepaticae)”. Here, Marchantiales is treated as circumscribed by
Parihar (1987) and Rashid (1998), and also as detailed earlier under the classification of Hepaticopsida
under Article 3.4.
Out of about 280 genera and approximately 9500 species of the class Hepaticopsida, Marchantiales
is represented by about 35 genera and approximately 420 species (Parihar, 1987). Marchantiales are
commonly called chambered hepatics.
COMMON GENERA 7.2

Besides Riccia and Marchantia, some of the other common genera of Marchantiales are Aitchinsoniella,
Asterella, Athalamia, Conocephalum, Cyathodium, Dumortiera, Mannia, Plagiochasma, Preissia,
Reboulia and Targionia.

Some of the salient features of Marchantiales are listed below:
1. Plant body is gametophytic, and gametophytes are usually prostrate, with dorsiventral thalli.
2. The thallus is usually dichotomously branched.
3. The dorsal surface is usually green, contains a midrib or mid-dorsal groove.
4. The ventral surface of thallus contains scales and rhizoids.
82 � Bryophyta
5. The rhizoids are of two types, smooth-walled and tuberculate.

6. The male and female sex organs (i.e. antheridia and archegonia) are either scattered along the
midrib, or they are grouped in usually raised receptacles.
7. Internally, the thallus shows well-marked specialisation of tissues in most Marchantiales. In a
few genera (e.g. Dumortiera), however, the differentiation of tissues is not well-marked.
8. The dorsal region contains several air chambers. Each air chamber opens outside by a well-
defined air pore.
9. The green tissue is mainly confined to the dorsal region of the thallus.
10. The ventral region of the thallus is usually made up of compact colourless parenchymatous
storage tissue.
11. Sometimes, some mucilage cells and cells filled with oil bodies are also present in the storage
region.
12. Gemmae, when present, are in a cup-shaped structure called gemma cup (e.g. Marchantia).
13. Archeogoniophore usually bears several archegonia, which ultimately develop into several
sporophytes.
14. The seta in the sporophyte is either short or absent.
15. The jacket or wall of the capsule is unistratose, i.e. single-layered in thickness.
16. In the sporogonia of Marchantiales, the columella is absent.
17. The capsule never opens by four regular valves. It dehisces in a variety of ways.
CLASSIFICATION 7.4
Campbell (1918) divided Marchantiales into three familes, viz. Ricciaceae, Corsiniaceae and
Marchantiaceae.
Verdoorn (1932) recognised six familes under Marchantiales. These are Marchantiaceae, Operculatae,
Astroporae, Targionaceae, Corsiniaceae and Ricciaceae.
Evans (1939) also recognised six families under Marchantiales, but these were not the same which
were recognised by Verdoorn. Six familes of Marchantiales recognised by Evans were Marchantiaceae,
Sauteriaceae, Rebouliaceae, Targionaceae, Corsiniaceae and Ricciaceae.
Campbell (1940) then revised his classification of Marchantiales on the basis of several new
features, including (i) nature and development of the receptacle, and (ii) structure of the sporophytes,
and divided this order into five families, viz. Ricciaceae, Corsiniaceae, Targionaceae, Monocleaceae
and Marchantiaceae.
Shiv Ram Kashyap, a well-known Indian bryologist, made detailed studies of liverworts of western
Himalayas and Punjab Plains and opined that all Marchantiales should be included only in three
families, namely Ricciaceae, Monocleaceae and Marchantiaceae.
In the later years, Carr (1956) and Proskauer (1961) also made detailed studies of several curious
thalloid hepatics (e.g. Carrpos sphaerocarus) and suggested several new changes in the classification
of Marchantiales.
Only Ricciaceae and Marchantiaceae are discussed here in some detail.
Hepaticopsida (Marchan ales) � 83
RICCIACEAE 7.5
A family of only three genera (Riccia, Ricciocarpos and Oxymitra) and about 150 species, Ricciaceae
members show the following general characteristics:
1. Ricciaceae are the simplest members of the order Marchantiales containing a gametophytic
thalloid plant body, which is usually prostrate, flat, dorsiventral and ribbonlike.
2. Upper or dorsal region of the thallus is called photosynthetic region, which contains either
large air chambers or narrow air canals enclosed by filaments of green, chlorophyll-containing
cells.
3. Well-marked pores are either absent or rudimentary.
4. A median longitudinal groove or strip, extending backwards from the growing apex, is present
on the dorsal surface.
5. Usually, the sex organs are borne in the region of the median longitudinal groove.
6. The sex organs (antheridia and archegonia) are usually immersed singly in cavities on the
dorsal surface of the thallus.
7. The neck and the apical portion of the archegonium usually remains projected above the
surface of the thallus.
8. The sporogonium remains usually sunken in the tissue of the thallus.
9. The sporogonium usually consists of only a saclike capsule. The foot and seta are absent.
10. The archesporium forms only the spores.
11. The elaters are absent.
12. The spores are disseminated or become free only when the thallus breaks down.
Life-history details of only Riccia are given here.
RICCIA 7.6
Order—Marchantiales
Family—Ricciaceae
Genus—Riccia

Riccia is a cosmopolitan genus of bryophytes. More than 200 of its species have so far been reported,
the majority of which are the inhabitants of the southern hemisphere. All the so-far reported species of
Riccia are terrestrial, except R. fluitans, which is the only aquatic, free-floating or submerged species.
The terrestrial species grow luxuriantly on damp soil and rocks. More than 30 species of Riccia have
so far been reported from India, specially from eastern Himalayas and the hills of southern India.
Some of the Indian species, growing commonly on moist soils, garden beds, stones and river banks are
Riccia discolor, R. cruciata, R. crystallina, R. pathankotensis, R. gangetica, R. billarderi, R. frostii and
R. melanospora (Fig. 7.1 A-F).
84 � Bryophyta
Fig. 7.1 A-F, External features of the gametophytes of some Indian species of Riccia. A, R. discolor
(R. himalayensis); B, R. billardieri; C, R. fluitans; D, R. pathankotensis; E, R. cruciata;
F, R. melanospora
Riccia fluitans, the only aquatic species, occurs floating in stagnant water or submerged a little
below the standing water surface.
The name Riccia has been given to the genus in honour of a Florentine politician, P F Ricci.
7.6.3 External Features of Gametophyte

The gametophytic plant body of Riccia is thalloid, flat, prostrate, fleshy and dorsiventral. The thallus
branches dichotomously, and due to repeated dichotomy, it becomes a rosette-shaped structure (Fig.
7.2 A). Each branch of the thallus is usually obcordate and sometimes linear to wedge-shaped and
ribbonlike.
A thickened mid-dorsal groove is present on the dorsal surface of the thallus (Fig. 7.2 B). This
groove extends up to the apical portion of the thallus, where it forms an apical notch. The apical notch
bears the growing point of the thallus.
The ventral surface (Fig. 7.2C) of the thallus possesses rhizoids and scales. The rhizoids are hairlike
elongated structures. Their main functions include (i) attachment of thallus to the substratum, and (ii)
absorption of water and soil solutes. Rhizoids of Riccia thus function as roots of higher plants. Rhizoids
are of two types, smooth-walled and tuberculate. The smooth-walled rhizoids (Fig. 7.2 D, E) are
unicellular, smooth-walled and elongated structures with colourless contents.
Fig. 7.2 External features, scales and rhizoids of Riccia. A, A rose e; B, Dorsal surface of thallus;
C, Ventral surface of thallus; D, Smooth-walled rhizoids in surface view; E, Smooth-walled
rhizoids in op cal sec on; F, Tuberculate rhizoids in surface view; G, Tuberculate rhizoids in
op cal sec on; H, A scale.
The tuberculate rhizoids (Fig. 7.2 F, G) have peglike or platelike ingrowths, which project into the
lumen from the wall. Rhizoids are absent in R. fluitans, the only aquatic species of the genus.
Scales (Fig. 7.2 H) are multicellular, one-cell-thick structures present on the ventral surface of the
thallus. These are violet-coloured bodies present on the margin. The violet colour is due to the presence
of a pigment, which remains dissolved in the cell sap. Scales are poorly developed or even absent
in some species, e.g. Riccia crystallina. The scales project forward near the growing point and thus
protect the young growing apex of the thallus.
Sex organs are present in the mid-dorsal groove region on the dorsal surface of the thallus. The
sporophytes, however, may be seen as black dots, when mature.
Riccia fluitans (Fig. 7.1C), the only aquatic spacies of Riccia, has a long, narrow, delicate, flattened,
ribbonlike, thalloid plant body. Its thalli are dichotomously branched. It lacks scales and rhizoids. In
the terrestrial conditions, the thalli of R. fluitans become thick, broadly channelled and also develop
numerous rhizoids as well as scales. Such scales are, however, confined to near the apex of the lobes
of the thallus.
7.6.4 Anatomy of the Thallus

The thallus appears like a boat-shaped structure in a vertical transverse section (Fig. 7.3 A), when viewed
under a dissecting microscope. It is thick in the midrib region and gradually becomes thin towards the
margins. The thallus is dorsiventrally differentiated into an outer or upper green photosynthetic region
86 � Bryophyta
and an inner or lower colourless storage region (Fig. 7.3 A, B). The storage region remains bounded
on the lower side by a layer of lower epidermis (Fig. 7.3 B), which bears two types of rhizoids in
the central region. These are smooth-walled rhizoids and tuberculate rhizoids. Compactly arranged,
parenchymatous, starch-containing cells are present in this ventral region, which is primarily a storage
tissue.
The upper, green photosynthetic region consists of more or loss erect vertical rows of unbranched
photosynthetic filaments or assimilatory filaments. These filaments remain separated by narrow air-
filled spaces called air chambers. Each cell of these filaments possesses numerous chloroplasts. Each
air chamber opens to the outside through a simple pore called air pore. The air pores are actually
the intercellular spaces between the upper epidermal cells (Fig. 7.3 B). The uppermost cells of the
photosynthetic filaments are comparatively large, lack chloroplasts and are thus colourless. These cells
form an ill-defined upper epidermis, the continuity of which is broken by air pores. The air chambers
communicate with the outer atmosphere on the dorsal surface of the thallus by air pores. Boat-shaped
vertical transverse sections of the thallus contain violet-coloured scales (Fig. 7.3 A), which are
multicellular structures, one cell in thickness.
In Riccia fluitans, the chlorophyllous tissue is made up of variously directed lamellae consisting of
one layer of cells (Fig. 7.3 C) enclosing large polyhedral air chambers. They are closed by an almost
continuous dorsal epidermis. Only a few air pores are present. Anatomy of some species is depicted
also in Fig. 7.3 D, E.
Fig. 7.3 Riccia. A, VTS of thallus (diagramma c); B, VTS of thallus (a part cellular);
C, VTS of thallus of Riccia fluitans; D, VTS thallus of R. glauca; E, VTS thallus of R. crystallina
7.6.5 Apical Growth

Thallus grows by the activity of a growing point located at the apex. This growing point consists of
three to five or more apical cells which are meristematic in nature. Each apical cell has four cutting
faces, of which one is dorsal, another is ventral and the remaining two are lateral.
In surface view, each apical cell appears rectangular, but in vertical longitudinal section, it appears
triangular (Fig. 7.4 A, B). Growth of the thallus is mainly contributed by segments which are cut
off from the dorsal and ventral faces of the apical cells. A major part of the thallus tissue is derived
from the dorsal segments. However, the lowermost cell layer of the thallus, rhizoids and scales are
contributed by the ventral segments.
Portions of the thallus contributed by five dorsal segments (D1, D2, D3, D4 and D5) and five ventral
segments (V1, V2, V3, V4 and V5) are shown in Fig. 7.4. Dichotomy in the thallus originated by
the cessation of the meristematic activity of one or more cells in a row of apical cells. Meristematic
activity of other adjacent apical cells results in the formation of two separate groups of cells, giving
rise ultimately to dichotomy.
Regarding the origin of the air chambers, some bryologists believed that the air spaces are depressions
in the surface of the thallus where growth of the tissue stops, and growth is more vigorous in the
adjacent parts, thus resulting in the formation of air chambers. This view has now been discarded.
Fig. 7.4 A, Ver cal longitudinal sec on through the growing point of thallus; B, Ver cal
longitudinal sec on of the thallus apex of Riccia glauca showing a triangular
apical cell and its deriva ves
Bryologists supporting the second view believe that air chambers develop “by splitting of the
internal cell walls in the originally compact upper tissue of the thallus, their origin being schizogenous
like that of the intercellular spaces in the parenchyma of vascular plants” as stated by Parihar (1987).
Such splitting of the internal cell walls begins endogenously, i.e. below and extends upwards (Pietsch,
1911). Some others, however, opine that splitting starts exogenously, i.e. begins at the surface and
extends downwards.

Some common methods of vegetative reproduction in Riccia are listed below:
1. Progressive Death and Decay This is the most common method of vegetative reproduction
in almost all species of Riccia. Progressive decay starts from the posterior part of the thallus, and
88 � Bryophyta
ultimately the older parts die. When this process of death and decay reaches up to the place of dichotomy,
the surviving branches of the thallus start behaving as independent thalli.
2. Adven ous Branches In species like Riccia fluitans, adventitious branches develop from
the ventral surface of the thallus. These branches, on separation from the main thallus, may develop
into new thalli.
3. Apex of Thallus In species, like R. himalayensis, the

apex of the thallus grows down into the moist soil and be-
comes thick at the end of the growing season. It develops
into a new thallus when favourable conditions return in the
next season.
4. Apex of Rhizoids In a few species, like R. glauca,

the apical part of the rhizoid has the ability to form a thallus,
in much the same way as a germinating spore forms a germ
tube and then a young thallus.
Fig. 7.5 Tubers of two species of Riccia.
5. Tubers Thalli sometimes develop perennating tubers, A, R. billardieri; B, R. discolor.
which have the ability to survive in unfavourable conditions.
On return of favourable conditions, these tubers may grow into new thalli, as in R. billardieri (Fig. 7.5
A), R. discolor (Fig. 7.5 B), R. perennis, etc.
7.6.7 Sex Organs

The two sex organs in Riccia are antheridia and archegonia. In monoecious or homothallic species (e.g.
R. billardieri, R. crystallina, R. gangetica, R. glauca and a majority of other species), both antheridia
and archegonia develop on the same thallus. But in dioecious or heterothallic species (e.g. R. bischoffi,
R. curtisii, R. discolor and some more species), antheridia and archegonia develop on separate thalli.
The sex organs develop in the region of median longitudinal furrow on the dorsal surface of the
thallus. This furrow extends backwards from the growing point. Sex organs develop in acropetal
succession, and serial sections through a thallus may reveal almost all stages of their development.
They arise from the segments which are cut off by an apical cell usually very close to, i.e. only 2-3 cells
away from the apical cell. Sex organs, in the different stages of their development, can be seen on the
same thallus. Due to the rapid development of the tissue surrounding the sex organs, they soon appear
to be embedded in the thallus (Fig. 7.3 D) or surrounded by a chamber-like structure called antheridial
chamber in case of male sex organ (i.e. antheridium) and archegonial chamber in case of female sex
organ (i.e. archegonium).
7.6.8 Development of Antheridium

Development of antheridium starts from a single superficial initial cell. This cell can be identified due
to its upward growth, and dense contents in relation to its neighbouring cells on the dorsal furrow, a few
cells behind the apical cells. It is known as antheridial initial (Fig. 7.6 A). Soon it becomes papillate
and divides transversely into a lower basal cell and an outer cell (Fig. 7.6 B). The basal cell remains
embedded in the thallus and forms the stalk of the antheridium. The outer cell projects slightly above
the thallus, undergoes several divisions and forms the rest of the antheridium. A series of transverse
divisions in the outer cell forms a filament of four superimposed cells (Fig. 7.6 C), of which two upper
cells are the primary antheridial cells and the two lower ones are the primary stalk cells. Two lower
primary stalk cells form the stalk of the antheridium, on which develops the globular body of the
antheridium. Two upper primary antheridial cells divide by two successive vertical divisions at right
Fig. 7.6 A-H, Development of antheridium in Riccia
angles to one another, and thus develop two tiers of four cells each (Fig. 7.6 D). Both the tiers of four
cells each now divide periclinally to from an outer layer of eight jacket initials, which are sterile, and a
central group of eight primary androgonial cells (Fig. 7.6 E, F), which are fertile. Jacket initials now
divide anticlinally to form a single layer of sterile jacket around the antheridium (Fig. 7.6 G). Repeated
divisions in the primary androgonial cells give rise to small cubical fertile androgonial cells, the last
generation of which are known as androcyte mother cells (Fig. 7.6 H).
Each androcyte mother cell divides diagonally into two triangular androcytes (Fig. 7.7 A). Each
androcyte is uninucleate and metamorphoses into an antherozoid (Fig. 7.7 B-F). During the process
of metamorphosis, each androcyte contains a prominent nucleus and an extranuclear granule called
blepharoplast (Fig. 7.7 B). The blepharoplast granule develops in the peripheral part of the protoplast
of androcyte. Soon the androcyte looses its triangular shape, and becomes a spherical body (Fig. 7.7 C).
The blepharoplast elongates in the form of a cordlike structure and occupies about 3/4th of the way of
the developing androcyte. The nucleus assumes a crescent shape, moves towards the periphery of the
protoplast and comes in contact with the blepharoplast (Fig. 7.7 D, E). Two flagella are produced from
one thickened end of the blepharoplast, which is quite a prominent structure (Fig. 7.7 F).
A mature antherozoid is uninucleate with a homogeneous nuclear portion, which occupies the
major part of the antherozoid. Blepharoplast, the extranuclear granule, ends in a head bearing two long
flagella (Fig. 7.7 F). The antherozoid moves with the help of its flagella. Both the flagella are attached
at the same level on opposite sides. The function of one flagellum is to help in propulsion while the
other flagellum helps in rotation and also in changing the direction of the antherozoid.
A mature antheridium (Fig. 7.6 H) consists of a small stalk and a globular or club-shaped body.
The stalk is short and few-celled and remains attached at the base of the antheridial chamber. The
90 � Bryophyta
body of the antheridium is composed of a central mass of either androcytes or antherozoids. It remains
surrounded by a single layer of sterile jacket, made up of tangentially elongated cells.
Antherozoids are liberated through a pore of the antheridial chamber, present on the dorsal surface
of the thallus. All cell walls within the mature antheridium disappear, and a semiliquid or viscous
content develops within. The antherozoids lie there in this viscous substance. Water drops enter within
the antheridial chamber. Cells of the antheridial jacket absorb this water by imbibition. They become
softened and finally break open to liberate the antherozoids, along with the semifluid mucilaginous
mass.
Fig. 7.7 A-F, Process of spermatogenesis and forma on of an antherozoid in Riccia
7.6.9 Development of Archegonium

Development of archegonium starts from a single superficial cell on the dorsal surface of the thallus,
quite close to the apical cell. This is called archegonial initial, which soon becomes papillate (Fig. 7.8
A) and divides by a transverse division into an outer cell and a basal cell (Fig. 7.8 B). The outer cell
ultimately forms the body of the archegonium while the basal cell finally develops into the embedded
portion of the archegonium. The outer cell divides by three successive intersecting vertical divisions to
form three peripheral initials and a fourth median cell called primary axial cell (Fig. 7.8 C-E).
Three peripheral initials divide by radial longitudinal walls to form six jacket initials, which
again divide by transverse walls to form two superimposed tiers of six cells each. Upper tier of six cells
form the neck and are called neck initials while the lower tier forms the venter, called venter initials.
Simultaneously, the primary axial cell divides transversely to form upper smaller primary cover cell
and the lower larger central cell (Fig. 7.8 F-I).
Repeated transverse divisions in the neck initials form a tube-like neck. It is made up of 6 to 9
cells in height and of six vertical rows of neck cells, as can be seen in the transverse section (Fig. 7.8
J). Simultaneously, six venter initials divide by several transverse and vertical divisions to form the
jacket of the venter. It is made up of 12 to 20 cells in perimeter.
Fig. 7.8 A-K, Showing development of archegonium in Riccia (Note: E, I and J are in transverse sec ons)
At the top of the neck is present the primary cover cell (Fig. 7.8 F). It divides by two successive
vertical divisions at right angles to each other and gives rise to four cover cells. The central cell (Fig.
7.8 F) divides transversely into an upper primary neck canal initial
and a primary venter initial (Fig. 7.8 G). The primary neck canal
initial divides transversely to form a row of four neck canal cells (Fig.
7.8 K). The primary venter initial also divides by one transverse
division to form an upper small ventral canal cell and a lower large
egg (Fig. 7.8 K).
A mature archegonium (Fig. 7.9) is a flask-shaped, long-necked
structure. It remains attached to the thallus tissue on the dorsal surface
by a short stalk. It contains a swollen venter and an elongated neck.
The neck is a single-layered, tube-like structure made up of 6 to 9
tiers of elongated cells arranged in six vertical rows and surrounding a
fine narrow canal. At the upper part of the neck are present four cover
cells. The canal of the neck contains four neck canal cells. The venter
contains a one-layered wall and encloses a small ventral canal cell
and a large egg. Immediately prior to fertilisation, the neck canal cells Fig. 7.9 A mature archegonium
and ventral canal cell disintegrate and form a mucilaginous mass. Due of Riccia.
to the pressure of this mucilaginous mass, all the four cover cells of the neck separate apart from one
another, and some of the mucilaginous mass also extrudes from the tip of the archegonial neck.
7.6.10 Fer liza on

The mucilaginous mass, formed by the disintegration of neck canal cells and venter canal cell imbibes
water, swells up and exerts a pressure on cover cells to separate. Due to all these activities, the canal
is open up to the venter. This mucilaginous mass also attracts spermatozoids swimming in the surface
film of water. The attraction is a chemotactic phenomenon due to the presence of some proteins and
92 � Bryophyta
inorganic salts in the mucilage. The spermatozoids are guided up to the egg due to this chemotactic
phenomenon. A spermatozoid (X) fuses with the egg (X), resulting into the formation of a diploid
structure called zygote (2X). The zygote soon secretes a wall, increases in volume and finally occupies
the entire lumen of the venter (Fig. 7.10 A–C).
7.6.11 Sporophyte
1. Development of Embryo The diploid zygote is the first cell of the sporophytic generation
or asexual generation. It soon secretes a cell wall of its own, enlarges and almost fills the cavity of the
venter (Fig. 7.10 C). Cells of the venter start dividing periclinally and then also anticlinally to form
a bilayered calyptra. Simultaneously, the zygote divides first by a transverse division (Fig. 7.10 D) to
form two almost equal-sized cells. Both these cells now divide vertically to form a four-celled embryo
or quadrant-stage (7.10 E). Soon follows one more vertical division at right angles to the first one and
thus results an eight-celled or octanct stage of the embryo. In this usual course of divisions resulting in
octanct stage of the embryo, there may be variations in different species of Riccia.
Fig. 7.10 A-J Some stages in the development of sporogonium, forma on of nutri ve fluid
and spore tetrads in Riccia
The young eight-celled embryo now divides in all the planes without any definite sequence to form
a multicellular (20 to 40-celled) spherical mass of cells (Fig. 7.10 F). It now divides periclinally into an
outer layer of amphithecium and an inner mass of cells, which represents the endothecium (Fig. 7.10
G). The amphithecium forms the jacket layer of the young sporogonium while the endothecium is
archesporial in origin. Cells of the jacket layer divide only by radial walls to form a single layer of
jacket. This layer is sterile and protects the sporogonium.
The endothecial archesporium, which is the first cell generation of sporogenous tissue, divides
and redivides (Fig. 7.10 H) several times to form a large mass of sporogenous cells. All these cells
are potential sporocytes or spore mother cells. Each of these spore mother cells is diploid, divides
reductionally and forms four haploid spores (Fig. 7.10 I, J). In Riccia crystallina and a few more
species, some spore mother cells fail to produce spores. Such spore mother cells form abortive nutritive
cells. Some bryologists (Pagan, 1932) opined that nutritive cells are forerunners of elaters, found in
some other Marchantiales (e.g. Marchantia).
The dividing spore mother cells are present in large amount of viscous nutritive fluid (Fig. 7.10 I),
which serves as nourishment for these spore mother cells and developing spores.
2. Sporogenesis Formation of haploid spores from the diploid spore mother cells is called sporo-
genesis. The sporogenesis starts with the contraction of cytoplasmic contents of spore mother cells or
sporocytes. Each spore mother cell divides by two successive divisions and develops into a tetrad of
spores (Fig. 7.10 J). It also results in a reduction division, thereby the chromosome number is reduced
to half, i.e. each diploid spore mother cell changes into four haploid spores arranged first in the form
of a tetrad. The cell plate is incomplete during the first division, and after the completion of the second
division, the cell plates delimiting four spores are formed simultaneously. All the four spores of a spore
tetrad remain opposed to one another and also remain surrounded by the wall of the spore mother cell
(Fig. 7.11 A-F) till the maturity of the spore wall.
3. Spore Each spore (Figs. 7.12 A-B; 7.13) is a uninucleate structure surrounded by three wall layers:
(a) The outermost perispore or exosporium is thin, mucilaginous but very strongly cutinized;
Fig. 7.11 A-F, Process of sporogenesis in Riccia. Fig. 7.12 Riccia spores. A, Surface view; B, Op cal view.
Fig. 7.13 A-E. A mature spore (A) and stages of its germina on and ini a on of a gametophyte (B–E) in Riccia
94 � Bryophyta
(b) Exine or mesosporium is a tough outer spore coat, in three concentric layers, and;
(c) Intine or endosporium is the innermost layer which is thin and homogenous.
The exine is variously ornamented. It shows reticulate sculptures, irregular reticulum or even
tubercles. Different species of Riccia may be identified on the basis of ornamentation patterns of exine.
In a majority of the species of Riccia, each spore of a tetrad has two faces, the convex distal face and
opposite to it are three flattened faces which form a pyramid-like structure.
4. Dispersal of Spore Spores in Riccia are dispersed without any definite mechanism, simply
(i) by disorganisation of wall of capsule or sporogonium, and (ii) by decaying of the surrounding tissue
of the thallus. Due to the absence of any definite mechanism, the spores may remain inside the thallus
for as long as one year or even more. Finally, they are dispersed by air currents or even by splashing
raindrops. On being dispersed, they germinate into gametophytes.
5. Germina on of Spore Spores start germinating only in moist conditions. At the time of
germination, a spore absorbs some water swells, and starts producing a germ tube (Fig. 7.13 A). Usu-
ally, the spores remain adhered in tetrads in the initial stages of their germination. The germ tube is an
elongated structure and remains filled with chloroplasts and oil globules. Its cell contents start accu-
mulating towards the apex. Soon the apical part of the germ tube is separated by a transverse division.
The so-formed cell now again divides by one more transverse division and then by a vertical division in
both these cells, resulting into a 4-celled body or germ disc (Fig. 7.13 B-C). From the lowermost part
of the germ tube also develops the first rhizoid (Fig. 7.13 B). Out of the four cells of the germ disc, one
starts functioning as an apical cell with two cutting faces. The activity of this apical cell results in the
formation of a multicellular young thallus (Fig. 7.13 D-E). It gets fixed in the soil by the development
of some more rhizoids from the newly-formed and young multicellular thallus. Sex organs develop on
such newly-formed thalli, and the life history (Fig. 7.14A-N) keeps on repeating again and again.
7.6.12 Life Cycle of Riccia

Diagrammatic life cycle is depicted in Fig. 7.14A–N.
MARCHANTIA 7.7
Order—Marchantiales
Family—Marchantiaceae
Genus—Marchantia

Marchantia, the best-known genus of the family Marchantiaceae, is represented by about 65 species,
distributed widely all over the world. The name to the genus was given in honour of Nicolas Marchant,
a French botanist. Chopra (1943) reported about 11 species of Marchantia from India, mainly from the
Fig. 7.14 A–N Diagramma c representa on of the life cycle of Riccia

Himalayas. All the so-far reported species are terrestrial and grow on the ground or soil in moist, shady
and cool conditions. It is also commonly seen on the sides of streams, wet rocks, damp burnt soil, walls
of the wells, swampy meadows and similar other surroundings.
96 � Bryophyta
Marchantia polymorpha is the best-known species. Kashyap (1919) reported M. palmata, M.

nepalensis and M. polymorpha from various parts of India. Udar (1970) reported 6 species from
different parts of India. The genus occurs commonly in almost all the hilly regions of India.
GAMETOPHYTIC PHASE
The gametophytic plant body is a thalloid, green, prostrate
and dichotomously branched structure with a dorsiventral
symmetry. Thalli attain a length of 2 to 10 cm or more.
Each lobe or branch of the thallus has a notch at the apex.
At the bottom of the notch is located the growing point.
On the dorsal or upper surface of the thallus (Fig.

7.15) is present a clear midrib. Several diamond-shaped
rhomboidal or polygonal areas are also present. The
boundaries between these areas mark the limit of the
underlying air chamber. A dotlike structure, present in the
centre of each polygonal area, represents the air pore.
Several thalli bear many cup-shaped, subsessile gemma
cups (Fig. 7.15) on the dorsal surface. Many gemmae
are present in each gemma cup. They are the means of Fig. 7.15 Marchan a polymorpha showing
vegetative reproduction. dorsal surface of the thallus
Mature thalli bear a few stalked, upright, fertile
branches, having either antheridia or archegonia at their
top portion. These antheridia-bearing branches are called
antheridiophores, while the archegonia-bearing branches
are called archegoniophores (Fig. 7.15).
On the ventral or lower surface of the thallus are present
scales and rhizoids (Fig 7.16).
Scales are violet to purple-coloured structures. They
are multicellular but one-celled thick structures. Scales are
arranged in 2 to 4 or more distinct rows on each side of
the midrib. Scales are of two types, i.e. appendiculate and Fig. 7.16 Marchan a nepatensis showing
ligulate. The appendiculate scales are situated usually ventral surface of the thallus
in one row just on both the sides of the midrib, and each
contains an appendage at its apical side (Fig. 7.17). The ligulate scales are present in one to three rows
on either side of the midrib. Each ligulate scale is ligule-like and is devoid of any appendage. The
ligulate scales are smaller than the appendiculate scales. The function of scales is to protect the growing
apex and to retain the moisture.
Rhizoids are unicellular, elongated and hairlike structures. They are of two types, i.e. smooth-
walled and tuberculate (Fig. 7.18). The smooth-walled rhizoids are broad, thin-walled, unicellular and
possess both the outer and inner smooth wall layers. The tuberculate rhizoids are comparatively narrow
and their inner wall layer contains many peglike ingrowths. Absorption and fixation are the functions
of the rhizoids.
Fig. 7.18 Rhizoids of Marchan a. A, Smooth-walled rhizoids in

surface view; B, Cross sec on of a smooth-walled rhizoid;
Fig. 7.17 An appendiculate scale C, Tuberculate rhizoids in surface view; D, Cross sec on of a
of Marchan a nepalensis tuberculate rhizoid; E, Both the rhizoids in cross sec on
7.7.4 Anatomy of the Gametophyte

Anatomically, the thallus is divisible into two different regions, i.e. upper photosynthetic region and
lower storage region (Fig 7.19, 7.20).
The photosynthetic region is
the green, dorsal or upper region
of the thallus, made up of upper
epidermis, air pores, air chambers
and photosynthetic filaments. The
upper epidermis is the outermost
layer made up of thin-walled cells.
Its continuity is broken by many
air pores. Each air pore is a barrel-
shaped, structure, usually surrounded
by 4-8 rings of superimposed cells
(Fig. 7.21 A-C). Each ring has 4-5 Fig. 7.19 VTS Thallus of Marchan a polymorpha (diagramma c)
cells. The wall of the pore lies half
above and half below the epidermis. Below the upper epidermis is present a single horizontal layer of
air chambers. The air chambers remain separated from each other by 2- to 6-celled partition walls.
These cells may or may not contain chloroplasts. Each air chamber opens outside by a barrel-shaped air
pore. Several green, chlorophyll-containing photosynthetic filaments are present in each air chamber.
These filaments consist of 2 to 6 or more cells and are branched or unbranched. The chlorophyll-
containing cells of these filaments constitute the main photosynthetic tissue of the gametophyte.
98 � Bryophyta
Fig. 7.20 VTS Thallus of Marchan a polymorpha (a part cellular)
Fig. 7.21 A, Air pore of Marchan a polymorpha (in surface view); B, Air pore of M. nepalensis in
ver cal cross sec on; C, Air pore of M. palmata (in surface view).
The storage region represents the ventral tissue of the thallus and lies below the photosynthetic
region. It consists of several layers of thin-walled parenchymatous cells. However, on the margins,
there are only 1 to 3 layers of this tissue. Usually, the cells are isodiametric in this region except along
the midrib. Most of the cells contain starch and are usually devoid of chloroplast. A few cells of this
region either contain a large oil body, or remain filled with mucilage. The cells of the midrib region
are marked with some reticulate thickenings. The cells of the lower epidermis contain both the types
of scales and rhizoids.

Marchantia reproduces vegetatively by a variety of ways mentioned below:
1. Death and Decay In this most common process of vegetative reproduction of thalloid bryo-
phytes, the basal or posterior part of the thallus starts rotting or disintegrating due to ageing or drought.
When this process of disintegration or decay reaches up to the place of dichotomy, the lobes of the
thallus get separated. All lobes, so detached, develop into independent plants by apical growth.
2. Adven ous Branches The adventitious branches may develop from the ventral surface or
from any other part of the thallus, and on getting detached, they develop into new thalli of Marchantia.
Kashyap (1919) reported the development of such branches from the stalk and disc of the archegonio-
phore in M. palmata.
3. Gemmae
Gemmae are the specialised means of vegetative reproduction in several species of Marchantia. They
develop in special cup-shaped structures, on the dorsal surface of the thallus, called gemma cups. Each
gemma cup (Fig. 7.15) is a circular or cup-shaped body with fringed margins, and usually develops in
the midrib region. Each gemma is a small, green, disclike and bilobed structure having a short, delicate
and unicellular stalk. Several small mucilaginous hairs also develop in the cavity of the gemma cup
along with the gemmae. Large amount of the mucilage is secreted by these mucilaginous hairs, which
finally help in the detachment of gemmae from the gemma cup.
(a) Development of Gemma Certain superficial initial cells, lining the floor of the gemma cup,
become active and appear as papillate outgrowths. These initial cells are called gemma initials. Each
gemma initial divides transversely into a lower basal cell and an upper cell. There is no further division
in the lower basal cell, and it forms the single-celled stalk. The upper cell divides transversely, and
both the so-formed cells undergo one more similar division resulting in a row of four cells. These four
cells divide several times in horizontal and vertical planes resulting in the formation of a thin plate-like,
multicellular gemma (Fig. 7.22 A-F). Several such gemmae, in different stages of development, are
seen in a gemma cup (Fig. 7.22 G).
(b) A Mature Gemma A mature gemma is a multicellular, lens-shaped structure, several cells thick in
the middle but only one-celled thick on the margins. It is deeply notched on the two opposite sides. A
growing point lies in each marginal notch. Each gemma contains a single-celled stalk. Majority of the
gemma cells contain abundant chloroplasts and are green. Some of its cells contain oil bodies instead
of chloroplasts. These are called oil cells. Some rhizoidal cells are also present on both the surfaces
of gemma.
Fig. 7.22 Marchan a. A-E, Stages of the development of a gemma; F, A mature gemma; G, Ver cal
cross sec on of thallus passing through gemma cup
100 � Bryophyta
(c) Germina on of Gemma The gemmae are dispersed by the wind or water current easily because of
their weak fragile single-celled stalk. When they fall on a suitable soil, the rhizoidal cells become active
and develop rhizoids. The meristematic cells, situated in both the marginal notches, start growing in
opposite directions and develop into two young thalli. The central portion of the parent gemma dies
leaving the production of two newly formed thalli of Marchantia.

The sexual reproduction is oogamous. The sex organs, i.e. antheridia and archegonia, develop on special
erect, stalked branches of the thallus. The antheridia-bearing erect branches are called antheridiophores,
and the archegonia-bearing branches are called archegoniophores. Marchantia is strictly dioecious.
The male thallus bears antheridiophores whereas the female thallus bears archegoniophores. The stalk
of both antheridiophores and archegoniophores contains a terminal horizontal disc. These erect sexual
branches are the direct continuations of the thallus, and this reflects also in their anatomical details.
7.7.7 Antheridiophore
1. Structure of Antheridiophore Antheridiophore has a long stalk of about 1-3 cm, bearing a
lobed disc at the apex. The disc is usually eight-lobed and rarely four-lobed. A growing point is situ-
ated at the tip of each lobe. Each lobe of the disc contains 2-7 or more antheridia arranged acropetally,
i.e. the oldest antheridium in the centre of the disc while the youngest antheridium near the apex of the
lobe. The antheridia are present on the dorsal surface of the disc. Each antheridium is enclosed in an
antheridial chamber.
2. Development of Antheridiophore It starts developing (Fig 7.23 A-F) as an outgrowth on

the male plant. Later on, the horizontal tubular outgrowth becomes erect and swells up at the tip. This
swollen tip develops into a lobed, concave, antheridial disc, bearing groups of antheridia on the dorsal
surface, and scales and rhizoids on the ventral surface.
Fig. 7.23 A-F, Development of antheridiophore in Marchan a polymorpha
3. Anatomy of the Stalk of Antheridiophore The stalk (Fig. 7.24) is more or less a pris-
matic, five-faced structure, of which three faces are furrowed and two faces are curved outward. Two
longitudinal furrows or grooves run down the length of the stalk. Scales and rhizoids are present in the
grooves, and thus the surface containing the grooves corresponds with the ventral surface of the thallus.
Several air pores, air chambers and photosynthetic filaments are present on the posterior side, which
thus corresponds with the dorsal surface of the thallus. The stalk thus shows a dorsiventral symmetry,
typical of the thallus.
Air chamber Air pore
Photosynthetic
filaments
Rhizoids
Groove
Fig. 7.24 T.S. Stalk of antheridiophore of Fig. 7.25 Longitudinal sec on (a part) of the
Marchan a polymorpha disc of antheridiophore of Marchan a
polymorpha
4. Anatomy of the Disc of Antheridiophore Anatomically, the disc also resembles the thal-
lus (Fig 7.25). The disc is surrounded by a layer of epidermis, the continuity of which is broken by
many barrel-shaped air pores. Each air pore opens in an air chamber having several photosynthetic
filaments. A growing region is situated at the apex of each lobe of the massive disc. Along with the air
chambers are also present several flask-shaped cavities called ‘antheridial chambers’. Each antheridial
chamber contains an antheridium and opens outside by a pore or ostiole. The antheridia remain ar-
ranged acropetally, i.e. the oldest antheridium is present in the centre and the youngest near the apex
of each lobe of the disc.
5. Development of the Antheridium The development of antheridium (Fig 7.26 A-I) starts
from a superficial antheridial initial cell on the dorsal surface of the disc of the antheridiophore (Fig
7.26 A). The antheridial initial enlarges in size, becomes papillate and divides first by a transverse
Fig. 7.26 A-I, Development of antheridium in Marchan a polymorpha

102 � Bryophyta
division forming an upper outer cell or primary antheridial cell and a lower basal cell (Fig. 7.26 B).
The outer cell or primary antheridial cell divides transversely into an upper antheridial cell and a lower
stalk initial cell (Fig. 7.26 C). The stalk initial cell divides by a few transverse and vertical divisions
and ultimately forms a small stalk of the antheridium. The upper antheridial cell divides transversely
to form a four-celled structure. This is followed by two vertical divisions at right angles to one another
forming a sixteen-celled structure, in which the cells are arranged in four tiers of four cells each (Fig.
7.26 D–E). A periclinal division in this sixteen-celled structure results in the formation of 16 outer
primary jacket cells and 16 inner primary androgonial cells (Fig. 7.26 F). The primary androgonial
cells divide by several repeated transverse and vertical divisions resulting in hundreds of androgonial
cells (Fig. 7.26 F-I). The primary jacket cells divide by several anticlinal divisions to form a single
layer of sterile antheridial jacket (Fig. 7.26 I).
Simultaneously, the cells, neighbouring the antheridial initial cell, also become active and develop
into a chamber-like structure around the antheridium. This chamber is called antheridial chamber.
6. Spermatogenesis The process of metamorphosis of androgonial cells into antherozoids is

called spermatogenesis. Each androgonial cell (Fig. 7.27 A) or androcyte mother cell is a cubic
structure and divides by a diagonal mitotic division to form two triangular androcytes. (Fig. 7.27 B).
Both the androcytes remain enclosed in the wall of the androcyte mother cell with no separating wall.
Each androcyte (Fig.7.27 C) has a prominent nucleus and a small extranuclear granule called
blepharoplast. The blepharoplast granule is present near the periphery of the protoplast. Soon the
androcyte loses its triangular shape and becomes somewhat round or oval. Its blepharoplast elongates
into a cord and occupies about two-third of its part. Simultaneously, the nucleus also becomes crescent-
Fig 7.27 A-G, Spermatogenesis in Marchan a polymorpha
shaped and homogeneous, and ultimately comes into contact with the blepharoplast. Two long flagella
are produced from the conspicuously thickened end of the blepharoplast. Thus, a uninucleate and
biflagellate antherozoid develops from each androcyte (Fig. 7.27 G).
6. Dehiscence of Antheridium Water helps in the dehiscence of antheridium. From the slightly
concave disc of antheridiophore, water enters into the antheridial chamber through its narrow canal.
When water comes into contact with some sterile cells of the jacket of the antheridium, they start disin-
tegrating. This results in the rupturing of antheridium. The androcytes in a group start emerging out of
the dehisced portion of antheridium in the form of a long smoke-like column. This mass of androcytes
breaks on reaching an air-water surface. Thus, the androcytes spread on the surface of water.
7. Mature Antheridium and Antherozoids A mature antheridium is a short-stalked, globular

and ovoid or club-shaped body present singly in an antheridial chamber. A single-layered sterile jacket
encloses the mass of androgonial cells.
Electron microscopic studies conducted by Sato (1951) indicate that the antherozoid of M.
polymorpha is a rod-like, uninucleate and biflagellate structure. Each flagellum is made up of several
fibrils. When the antherozoids swim in water, they resemble the crawl of a snake.
7.7.8 Archegoniophore
1. Structure and Development of Archegoniophore Similar to antheridiophore, the arche-
goniophore or carpocephalum also consists of a stalk and a disc. The stalk is simply the modified
branch of the thallus. The disc is usually eight-lobed. In M. polymorpha, the stalk of archegoniophore
terminates in a nine-rayed stellate disc. The stalk of the archegoniophore is comparatively smaller than
the stalk of antheridiophore.
When very young, the apex or growing point of the archegoniophore protuberance becomes swollen
(Fig. 7.28 A, B). The dichotomy is repeated in quick succession in this swollen apex, which ultimately
becomes a rosette-like, eight-lobed disc. From the original branch initial, an eight-lobed disc develops
by three successive dichotomies. Each lobe of the disc contains a growing point. Soon a group of
archegonia develops in each lobe in acropetal succession, i.e. youngest near the apex and the oldest
archegonium in the centre of the disc. Thus, eight groups of archegonia are seen on the upper surface
of the disc.
Fig 7.28 A-F, Development of archegoniophore in Marchan a polymorpha
The archegonia develop in erect position on the archegonial lobe with their necks directed upwards
(Fig. 7.28 C). The erect position of the archegonia is seen only when the stalk of the archegoniophore is
very short and the disc is only slightly raised above the thallus. Fertilization takes place at this stage.
Immediately after fertilisation, following changes occur in quick succession:
(a) Stalk of the archegoniophore begins to elongate with the simultaneous overgrowth in the
central sterile part of the disc.
(b) Because of the overgrowth of the central part of the disc, the groups of archegonia and the
marginal apical regions of the disc are shifted towards the lower side (Fig. 7.28 D, E). This
brings the curvature of the apices of lobes.
(c) The growing apex of each lobe of the disc now lies near the stalk of the archegoniophore.
(d) The archegonia are now hanging towards the lower side with their necks pointing
downwards.
(e) Because of the curvature of the apices, the youngest archegonium is now present near the
stalk of the archegoniophore while the oldest archegonium near the periphery of the disc
(Fig 7.29).
104 � Bryophyta
Air chamber
Archegonia
Perigynium
Air pore
Rhizoids Young
sporophyte
Perichaetium
Ray
Stalk
Fig 7.29 LS (a part) of archegoniophore disc a er fer liza on, showing the youngest
archegonium near the stalk
(f) At the base of each archegonium develops a ring of cells in the form of a collar-like structure
called perigynium or pseudoperianth.
(g) A bilipped, pendant, one-celled thick sheath, with fringed margins, develops from both the
sides of the group of archegonia. This involucral sheath encloses the group of archegonia and
is known as perichaetium (Fig. 7.29).
(h) Between the groups of archegonia, in some species, develop long, green, cylindrical processes
from the peripheral portion of the disc. These processes are called rays. The rays provide an
umbrella-shaped structure to the disc. In M. polymorpha, the rays are usually nine in number.
2. Anatomy of Archegoniophore Anatomically, the stalk of an archegoniophore resembles the

stalk of an antheridiophore (Fig. 7.24). Its morphologically ventral side has two longitudinal grooves
having scales and rhizoids, whereas its morphologically dorsal side, resembling the dorsal surface of
the thallus, contains air chambers, air pores and photosynthetic filaments. In the upper side of the disc
portion are also present the air chambers, air pores and photosynthetic filaments (Fig. 7.29), Groups of
archegonia, perigynium, perichaetium and rays are also present in the disc region of the archegonio-
phore.
3. Development of Archegonium The development of archegonium starts from a dorsal su-

perficial cell which acts as an archegonial initial (Fig. 7.30 A). The archegonial initial enlarges, be-
comes papillate and first divides transversely into a lower basal cell and an upper outer cell (Fig 7.30
B). There is no further division in the basal cell, and thus the entire archegonium develops from the
outer cell.
The outer cell divides by three successive intersecting walls or periclinal vertical divisions resulting
in the formation of three outer peripheral cells and a central axial cell (Fig. 7.30 C-H). The axial cell
divides transversely and unequally, forming a small upper cover initial cell and a large central cell
(Fig.7.30 G). Each of the three peripheral cells divides by an anticlinal vertical division forming two
cells. In this way, the axial cell gets surrounded by six peripheral cells (Fig. 7.30 J). All the peripheral
cells divide transversely into upper neck initial tier and lower venter initial tier. Simultaneously, the
large central cell also divides transversely into an upper neck canal initial and a lower central cell
(Fig. 7.30 I). The primary neck canal initial divides by a series of transverse divisions to form 4 to 6
neck canal cells. The central cell divides transversely only once and forms a small venter canal cell
and a large egg (Fig. 7.30 J).
The cover initial cell divides by two vertical divisions at right angles to one another forming four
cover cells which form the mouth of the archegonial neck. The cells of the neck initial tier divide by
repeated transverse divisions to form a long neck. The cells of the venter initial tier also divide by
repeated transverse divisions to form a single-layered swollen venter (Fig. 7.30 K,L).
4. Mature Archegonium A mature archegonium (Fig. 7.30 L) is a flask-shaped structure with a long
neck and a globular venter. The neck consists of six vertical rows of cells enclosing 4 to 6 neck canal
cells. The venter consists of a single-layered jacket enclosing a venter canal cell and an egg. Four cover
cells form the mouth of the archegonium.
Fig. 7.30 A-L. Development of archegonium in Marchan a polymorpha

106 � Bryophyta
7.7.9 Fer liza on

Fertilization takes place at a time when the archegoniophores are only slightly elevated above the
thallus, and the necks of the archegonia are facing upwards. Water is essential for fertilization. A
mass of antherozoids exudes from the canal of the antheridial chamber. Antherozoids keep freely
swimming in the water. The antherozoids are positively chemotactic to certain substances, such as
proteins, inorganic salts of potassium, etc. An antherozoid enters the archegonial neck because of the
chemotactic response. Two gametes, i.e. antherozoid and egg, fuse, but, according to Anderson (1929),
the nuclei of the two gametes fuse only after a long time. The fusion of the nuclei of male and female
gametes results in the formation of a diploid structure called zygote.
SPOROPHYTIC PHASE
7.7.10 Post-fer liza on Changes
After fertilization, the oospore or fertilized egg, enlarges until it completely fills the cavity of the
venter of the archegonium. A cellulose cell wall is then secreted around the oospore. The surrounding
gametophytic tissue is also affected due to the developing oospore, which is the first cell of the
sporophytic generation. A few noticeable changes take place in the developing sporogonium and the
surrounding tissues. These are mentioned below:
1. The stalk of the archegoniophore starts elongating and becomes 3 to 4 cm long.
2. Periclinal divisions take place in the cells of the venter. This makes the venter two- to three-
layered, and it is’now called calyptra.
3. The calyptra later on expands and protects the young developing sporogonium.
4. Several cells, in the form of a ring, at the base of the venter become active, divide and redivide
and form a thick, collar-like outgrowth called perigynium or pseudoparianth. When young,
the perigynium is only one-cell thick. It forms a close covering at the base of the archegonium
enclosing the young sporogonium. Its function is to provide protection to the developing
sporogonium.
5. The group of archegonia, a few of which contain the eggs and a few others the developing
sporogonia, is also covered by yet another protective covering called perichaetium.
The young sporogonium is thus surrounded by three protective coverings of gametophytic origin,
i.e. calyptra, perigynium and perichaetium.
7.7.11 Development of Sporogonium

The oospore or zygote (Fig. 7.31 A) first divides by a transverse division to form an upper epibasal
cell and a lower hypobasal cell (Fig. 7.31 B). Further development of the young embryo is different in
different species. In several species, including M. polymorpha, the second division is at right angles to
the first one to form a 4-celled quadrant stage (Fig. 7.31 C). The epibasal derivatives of this quadrant
stage form the capsule while the hypobasal ones form the seta and foot.
In M. domingensis, the quadrant stage develops as in M. polymorpha, but the epibasal part of the
quadrant develops into a capsule and a part of seta while the hypobasal part gives rise to the remaining
basal part of seta and capsule.
In M. chenopoda, the second transverse division is parallel to the first transverse division resulting
in a 3-celled filamentous embryo, of which the uppermost cell gives rise to the capsule, the middle cell
to seta and the lowermost cell to foot.
In the 4-celled stage of M. polymorpha, the next division is also a vertical division but at right angles
to the first one to form an 8-celled stage of the embryo called octant stage. The young embryo now
begins to elongate at this stage. There are now differences in the subsequent divisions in both hypobasal
and epibasal regions. Four cells of the hypobasal region of the octant stage of the embryo divide and
redivide to form a parenchymatous tissue (Fig. 7.31 D). From the lower cells of this parenchymatous
tissue develops the foot while from its upper cells develop the seta.
Foot is the lowermost region of the sporogonium. It is made up of a bulbous mass of cells, which
presses itself into the gametophytic tissue. It functions as an anchoring and absorbing organ of the
sporogonium.
Seta is the middle region of the sporogonium. It arises from the upper cells of the parenchymatous
tissue which develop from the hypobasal region of the embryo in M. polymorpha. The cells in the seta
are arranged in regular vertical rows. When young, the cells of the seta region are isodiametric in shape,
but, they soon divide by repeated transverse divisions which ultimately result in an increase in the
seta length. The elongating seta pushes the capsule, which ruptures the calyptra and other membranes
surrounding the sporogonium (Fig 7.31 E-G).
Fig. 7.31 A–G, Development of sporogonium in Marchan a polymorpha; H, L.S. of a mature

sporogonium; I, spore mother cells and elater mother cell; J, spore tetrad; K, Two elaters.
108 � Bryophyta
Capsule is the uppermost part of the sporogonium. It develops from the four cells of the epibasal
region of the octant stage of embryo in M. polymorpha. The cells of this region divide irregularly to
form a round mass of cells. With the help of a periclinal division, in the cells of the peripheral region
of this mass of cells, two layers are formed, an outer layer of amphithecium and the inner cellular
mass called endothecium. The cells of the amphithecial layer divide only by anticlinal walls and form
a single-layered jacket or capsule wall. Some annular thickenings may also develop in the cells of the
capsule wall in the later stages.
The archesporium is endothecial in origin in Marchantia. Massive sporogenous tissue develops
by several divisions in the cells of the endothecium. About half of the cells of the sporogenous tissue
divide by several transverse divisions to form fertile spore mother cells or sporocytes. These spore
mother cells are more or less cubic in shape and arranged in the form of vertical rows. According to
O’Hanlon (1926), each sporogenous cell in M. polymorpha divides by five successive divisions to form
32 spore mother cells. However, only 8 or 16 spore mother cells are formed by three or four successive
divisions of a sporogenous cell in M. domingensis, as reported by Andersen (1929).
The remaining half of the cells of the sporogenous tissue become sterile and form elaters. The
elaters (Fig. 7.31 K) are long, spindle-shaped, slender cells possessing tapering ends. On the inner
surface of the cell walls of each of these elaters are present two spiral thickenings, formed by the
partly disappearance of their protoplasm. The elaters are hygroscopic in nature and help in loosening
of spore mass and dispersal of spores. In M. polymorpha, an elater is equivalent to a sporogenous cell.
As mentioned above, a fertile sporogenous cell produces 32 spore mother cells by five successive
divisions. Thus, there are 32 spore mother cells (or 128 spores) to each elater in the sporogonium of
M. polymorpha.
Each spore mother cell is diploid and divides meiotically to form four haploid spores (Fig. 7.31 I, J),
which remain arranged tetrahedrally for quite some time (Fig. 7.31 J). The spores later become free and
remain enclosed by the capsule wall along with elaters. It has been estimated that as many as 300,000
spores may be produced in a single sporogonium of
M. polymorpha (O’Hanlon, 1926).

A mature sporogonium of Marchantia is an elongated
structure differentiating into three regions, i.e. foot,
seta and capsule (Figs 7.31 H, 7.32). The foot is
multicellular, bulbous, anchors the sporogonium to
the disc of the archegoniophore, and also helps in the
absorption of food for the developing sporogonium
from the gametophyte. The seta is a thick, stalk-like
structure in between the foot and capsule. It consists
of cubic cells arranged in vertical rows, and its
function is elongation. The capsule is the uppermost,
spherical or oval, and spore-producing region of the
sporogonium. The capsule is surrounded by an outer
layer of sterile wall called jacket or amphithecium.
The cells of the wall have ringlike thickened bands. Fig. 7.32 L.S. mature sporogonium of Marchan a
polymorpha
Inside the wall are present the fertile spore mother cells and sterile elaters. Elaters are long, spirally
thickened structures with tapering ends. The spore mother cells divide reductionally to form haploid
spores. Ruptured calyptra, perigynium and perichaetium coverings are also present around the mature
sporogonium.
7.7.13 Dehiscence of Capsule

After the formation of haploid spores from the diploid spore mother cells in the capsule, the seta
elongates very fast. It pushes the capsule out of the protective coverings of calyptra, perigynium and
perichaetium. The exposed capsule starts drying from the top. The long elaters start twisting and create
tension inside the capsule. Due to this process, the jacket of the capsule splits longitudinally, from the
apex to about the middle, into a variable number of valves. The hygroscopic nature of elaters and cells
of the capsule jacket also helps in the splitting of the capsule. The jerking action of elaters also helps in
loosening the spore mass. Wind helps in the spore dispersal from the dehisced capsule.
YOUNG GAMETOPHYTE
7.7.14 The Spore
The spores are very small (from 0.012 to 0.03 mm in diameter), haploid, round or spherical, uninucleate
structures surrounded by two layers, i.e. an outer thick, smooth or ornamented exine, and an inner
thin and smooth intine. Granular cytoplasm surrounds the nucleus. The spores in species, such as M.
polymorpha and M. faleacea are apolar, i.e. they lack a definite polar axis and are globose in outline,
while in species, such as M. tonasa, the spores are cryptopolar, i.e. they possess a polar axis and are
not globose in outline.
The spores are of about 10-12 m in diameter in M. polymorpha, 18-25 m in diameter in M. nepalensis,
and 24-30 m in diameter in M. palmata.
7.7.15 Germina on of Spore and Forma on of Young Gametophyte

The spore germination (Fig 7.33 A-H) starts immediately if favourable conditions persist. The viability
of spores is almost 100% for one full year. At the time of germination, the spore first swells and
becomes nearly double the original size. Out of the four spores of a tetrad, usually two develop into
male thalli and the remaining two into female thalli.
The early divisions in the germinating spore are in one plane only, and thus develop short
filamentous structures. In the later stages, however, the divisions are in all planes. When about a
dozen cells are formed in the young gametophyte of M. polymorpha, a marginal row of cells become
conspicuous according to O’Hanlon (1926). This row of cells is responsible for further growth of the
young gametophyte. A notch develops in the apical region when the gametophyte contains about 25-40
cells. However, Menge (1930) opined that when the young gametophyte is only 6- to 8-celled in M.
polymorpha, there develops an apical cell with two cutting faces. This apical cell cuts cells in both left
and right sides by repeated divisions, and thus develops a young gametophytic thallus of Marchantia.
110 � Bryophyta
Fig. 7.33 A–H, Spore germina on in Marchan a polymorpha.
7.7.16 Life Cycle of Marchan a

See Fig 7.34.
Fig. 7.34 Diagramma c life cycle of Marchan a

COMPARISON OF SPOROGONIUM OF RICCIA AND

MARCHANTIA 7.8
Table 7.1 Comparison of sporogonium of Riccia and Marchan a
S. NO. RICCIA SPOROGONIUM MARCHANTIA SPOROGONIUM

1. Sporogonium is simplest amongst bryophytes. Sporogonium is a very complex structure.
2. It is made up of only capsule or spores. It lacks It is made up of well-developed foot, seta and
foot and seta. capsule.
3. The mature sporogonium remains almost sunken It remains attached to the thallus only by foot.
in the thallus. It hangs from the undersurface of the female
receptacle.
4. The sporogonium remains surrounded by a Only the capsule remains surrounded by a single-
jacket layer made up of sterile cells. layered capsular wall.
5. Jacket layer surrounds the fertile spore mother Jacket layer of capsule surrounds the mass of
cells. There are no elater mother cells. fertile spore mother cells and sterile elater mother
cells.
6. Few sterile nurse cells are present in the Nurse cells are absent. The sterile tissue of the
sporogonium. sporophyte is in the form of foot, seta, capsule
wall and elater mother cells.
7. Elaters are absent. Elaters are present.
8. Mature sporogonium has almost no diploid tis- Mature sporogonium completely made up of
sue. The outer layer of sporogonium represents diploid tissues, except spores which represent the
parent gametophyte and spores represent future new gametophyte.
gametophyte.
9. A bilayered calyptra surrounds the sporogo- A bilayered calyptra, and well-developed cover-
nium. The layers like perigynium and per- ings or layers of perigynium and perichaetium are
ichaetium are absent. present.
10. Dehiscence of capsule never occurs. The disper- Dehiscence of capsule takes place by 4-6 valves,
sal of spores is simply by the disintegration of and spores are dispersed into the air.
the capsule wall.
11. Mature haploid spores are simply left behind Mature haploid spores are dispersed by the wind
on the soil due to the decay of the surrounding under a specialised process.
thallus tissue.
ORIGIN AND EVOLUTION OF MARCHANTIACEOUS THALLUS 7.9

Bryologists may be grouped in two different categories in suggesting their theories regarding the origin
and evolution of Marchantiaceous thallus. Some believe that the simple looking thallus of members
of Marchantiales is due to retrogressive evolution while other bryologists believe that simplicity of
thallus of these members is the result of progressive evolution. Let us explain both these theories as
under.
112 � Bryophyta
7.9.1 Retrogressive Evolu on

Bryologists believing retrogressive evolution of Marchantiaceous thallus may also be placed in two
different groups. Majority of them believe that retrogressive evolution has taken place by progressive
simplification, called reduction. On the other hand, some believe that it has been brought about by
condensation. The former view has been termed reduction theory, while the latter view has been
expressed under condensation theory.
1. Reduc on Theory
(a) Theory of Von We stein Von Wattstein (1908) was the first to propose the reduction theory
of retrogressive evolution of origin of Marchantiaceous thallus, and received the support of several
top bryologists of those days, including Church (1919), Kashyap (1919), Goebel (1930) and Evans
(1939). In his theory, Von Wettstein opined that “the primitive gametophyte of Hepaticae was nearest
to the erect, leafy Acrogynous Jungermanniaceous forms”. He treated Acrogynous Jungermanniales
of the Calobryum-type first. Plant body in Calobryum is an erect, leafy gametophyte showing radial
symmetry and containing leaves in three rows. During the course of reduction, the first step was the
adoption of a prostrate habit. Because of the development of dorsiventral habit, the leaves on the
ventral side gradually reduced in size, and finally in some cases, they disappeared completely. These
changes in habit and reduction of size of ventral leaves were soon accompanied by the flattening of
central axis of the gametophyte. The lateral leaves were first partially and then completely eliminated.
The final result of all these changes was the “formation of a leafless, flat, dorsiventral Acrogynous
Jungermanniaceous gametophyte of Pellia-type” (Von Wettstein, 1908). In the later stages, there was
“gradual progressive internal differentiation of tissues”, which finally resulted into the “formation of
externally simple but internally highly differentiated thallus of Marchantiales” (Von Wettstein, 1908).
He believed that scales on the ventral surface of the Marchantiaceous thallus as ‘modifications of the
leaves of the ventral row of the prostrate, foliose ancestor’.
(b) Kashyap’s Reduc on Theory of Pteridophytean Origin of Marchan ales SR Kashyap (1919)
believed in the reduction theory but opined differently from that of the theory of Von Wettstein.
He postulated the theory of Pteridophytean origin and opined that liverwort thallus, in general, and
Marchantiaceous thallus, in particular, shows great resemblance in external form and structure with
the prothalli of Lycopodium cernuum and Equisetum debile of pteridophytes. He also correlated the
erect, chlorophyll-containing branched lobes of dorsal photosynthetic region of L. cernuum and E.
debile with the erect photosynthetic filaments and walls of air chambers of Marchantiales. On the basis
of these similarities between some pteridophytes and Marchantiales, Kashyap (1919) suggested that
“Marchantiales probably arose by reduction from these pteridophytean ancestors”.
Kashyap’s reduction theory of pteridophytean origin of Marchantiales was, however, opposed by
P N Mehra, another Indian bryologist, on the basis of the following arguments:
(i) Pteridophytes have a well-developed vascular system, while liverworts, in general, and
Marchantiales, in particular, have no evidence of any lost vascular system.
(ii) Simple sporophytes of Marchantiales have many clear dissimilarities with the sporophytes of
Lycopodium and Equisetum.
(iii) Scales, present on the ventral surface of the thallus of Marchantiales, have no similarity with
any structure of the undersurface of the prothallus of Lycopodium cernuum and Equisetum
debile.
(iv) There exists no similarity between the sex organ-bearing erect structures (antheridiophores and
archegoniophores) of Marchantia and sex organ-bearing parts of Lycopodium and Equisetum.
2. Condensa on Theory
Mehra and Vasisht (1950) proposed that the thalloid forms of Marchantiales and other liverworts
have been derived from the foliose forms by the process of condensation, and this theory is known as
condensation theory. On the basis of these findings in the later years, Mehra (1957) observed the real
phylogenetic connection between thalloid forms of Marchantiales and foliose forms of Jungermanniales
and explained the origin of Marchantiaceous thallus by compaction, condensation and fusion of leaves
of Jungermanniales. Major outlines of the condensation theory are listed below:
(a) The lateral leaves of the foliose ancestors overlapped succubously or incubously.
(b) The lower portions of these leaves fused at the points of contact. This resulted in the formation
of single-layered wings on either side of the central axis. This also resulted in the formation of
lamellae on the upper side of wings.
(c) Thus formed lamellae were one-cell thick.
(d) These lamellae were directly obliquely outwards and were responsible for the formation of
open chambers.
(e) Then there was the inward growth of the margins of the cavities of these open chambers, and
this resulted in the formation of roof of these air chambers.
(f) In the early stages, the communication of these roofed chambers with the exterior area was by
large gaps.
(g) These gaps were later on replaced by air pores.
(h) Development of the air pores was also helpful in protection against loss of water by
transpiration process.
(i) Simultaneously, there was a gradual flattening of central axis.
(j) Flattening of the central axis reached up to the margins of the wings.
(k) Along with the flattening process, there occurred the development of secondary partitions
across the air chambers.
(l) Horizontal partitions also developed between the lamellae in xerophytic conditions, resulting
in the formation of several-layered chambers, as in Plagiochasma.
(m) Along with all the above-mentioned changes, there developed photosynthetic filaments from
the floor of air chambers, as in Marchantia.
(n) In Marchantia and a few other Marchantiales, there also developed barrel-shaped air pores
in air chambers.
7.9.2 Progressive Evolu on

Originally proposed by Schiffner (1893), the theory of progressive evolution was supported by some
famous bryologists, such as F Cavers (1903), FO Bower(1935), DH Campbell (1936), GM Smith (1955)
and others. The progressive evolution theory suggests that the Hepaticae members “originated from a
simple thallose gametophyte”. The ancestral thallus was simple, prostrate, and exhibited no external
or internal differentiation. According to Cavers (1903), the forms like Sphaerocarpos, are the present-
day Hepaticae, while Campbell (1936) opined that forms like Metzgeria are the present-day Hepaticae
which exhibited a nearest approach to the primitive Hepaticean gametophyte. The following two types
of gametophytes evolved from these primitive types by progressive evolution process:
114 � Bryophyta
1. Marchan aceous Gametophytes Marchantiaceous gametophytes evolved from the primi-

tive thallose gametophyte by progressive evolution as under:
(a) Simple, prostrate, thallus-like form of primitive thalloid gametophyte has been retained as
such.
(b) During evolution, there has been a gradual but progressive internal differentiation of tissues.
Finally developed the thallus, made up of (i) a definite epidermis, (ii) well-developed air pores
opening into the air chambers, (iii) air chambers containing photosynthetic filaments, and
(iv) well-organised parenchymatous and compact region representing the ventral surface or
storage tissue of the thallus.
(c) Sex organs aggregated into specified areas called receptacles, e.g. antheridia on antheridiophore,
and archegonia on archegoniophores.
2. Jungermanniaceous Gametophytes Jugermanniaceous gametophytes evolved from the

primitive thallose gametophyte by progressive evolution as under:
(i) Simple internal structure of primitive thalloid gametophyte has been retained.
(ii) There has been a gradual but progressive elaboration of the external form.
The above-mentioned changes resulted in the evolution of a leafy, prostrate thallus of members of
Jungermanniales.
1. What do you mean by Marchan ophyta?

2. Marchan ales are commonly called _____.
3. Two major genera of Marchan ales are Marchan a and _____.
4. Name any five common genera of Marchan ales.
5. Enlist any seven general characteris cs of Marchan ales.
6. Ventral surface of the thallus of Marchan ales contain rhizoids and _____.
7. Two types of rhizoids of Marchan ales are smooth-walled and _____.
8. In Dumor era, the differen a on of ssues is _____ well-marked
9. In the thalli of Marchan ales, the green ssue is mainly confined to which region: dorsal or
ventral?
10. Usually, the jacket wall of the capsule of Marchan ales contains how many layers?
11. In the sporogonia of Marchan ales, the columella is _____
12. Write a note on classifica on of Marchan ales in about 100 words.
13. Three genera of Ricciaceae are Riccia, Ricciocarpos and ____.
14. Write any five general characteris cs of the family Ricciaceae.
15. Name the aqua c species of Riccia.
16. Are a majority of the species of Riccia aqua c or terrestrial?
17. Write a brief life-history account of Riccia in about 1000 words
18. Give in detail an illustrated account of the external features of the gametophyte of Riccia.
19. Write short notes on the following:
(a) Scales of Riccia (b) Rhizoids of Riccia
(c) Anatomy of the thallus of Riccia fluitans (d) Apical growth in Riccia
20. How does Riccia reproduce vegeta vely?

21. Discuss in brief the development of sex organs in Riccia. Support your discussion with suitable
diagrams.
22. Give an account of spermatogenesis and forma on of an antherozoid in Riccia
23. How many flagella does a mature antherozoid of Riccia contain?
24. Write illustrated notes of the structure of mature sex organs of Riccia.
25. A mature archegonium of Riccia is _____ shaped.
26. Discuss in detail the sporogonium and its various stages of development in Riccia.
27. Write a note on nutri ve cells of the sporogonium of Riccia.
28. Draw diagramma c representa on of the life-cycle of Riccia.
29. Name the best-known species of Marchan a.
30. Explain in detail the external features of the gametophyte of Marchan a.
31. Write a detailed note on the gemma cups in Marchan a.
32. Write one major difference between the scales of Riccia and Marchan a.
33. What are the major differences between the scales of Riccia and Marchan a?
34. Discuss the anatomy of the Marchan a thallus.
35. Air pores of Marchan a are _____ shaped.
36. What are gemmae? Explain their development in Marchan a.
37. Write major details of the life history of Marchan a in about 1000 words.
38. Write detail illustrated notes on:
(a) Antheridiophore of Marchan a
(b) Archegoniophore of Marchan a
39. Explain in detail the sporophyte and its development in Marchan a.
40. In the sporophyte of Marchan a, what are calyptra, perigynium and perichae um?
41. Marchan a sporophyte is divisible into foot, _____ and capsule
42. Make a well-labelled diagram of LS of the mature sporogonium of Marchan a.
43. Compare the sporogonia of Riccia and Marchan a.
44. Explain in detail the origin and evolu on of the Marchan aceous thallus.
8
Anthoceropsida
WHAT IS ANTHOCEROPSIDA? 8.1

The class Anthoceropsida (Rothmaler, 1951), variously named as ‘Anthocerotes’ (Howe, 1899; Campbell,
1918), or ‘Anthocerotae’ (Smith, 1938, 1955; Takhtajan, 1953; Wardlaw, 1955) or Anthocerotopsida
(Proskauer, 1957), is a small group of bryophytes, and its best known genus is Anthoceros. The members
of this class are commonly known as horned liverworts.

Some of the generalized characteristics are described here:
1. The plant body is gametophytic and the gametophytes are dorsiventral, thalloid and variously
lobed.
2. The thallus does not show any internal differentiation of tissues.
3. On the ventral surface of the thallus are present smooth-walled rhizoids. The tuberculate
rhizoids, found in several members of Hepaticopsida (e.g. Riccia, Marchantia), are absent.
4. The scales are absent.
5. The air chambers and the air pores are absent.
6. Each cell of the thallus contains one or sometimes more, large, laminate chloroplasts, and each
chloroplast contains a pyrenoid.
7. Oil bodies are absent in the cells.
8. Antheridia develop from the hypodermal cells on the dorsal side of the gametophyte.
9. Antheridia are present either singly or in groups, in closed cavities called antheridial
chambers.
10. Archegonia are almost completely embedded in the gametophytic tissues on the dorsal surface
of the thallus.
11. The sporophyte consists of a bulbous foot, an intercalary meristematic region instead of seta,
and a long capsule.
Anthoceropsida � 117
12. Owing to the presence of the intercalary meristem, the sporophytes continue to grow
indefinitely, i.e. show indeterminate growth.
13. A well-developed central columella is present in each sporophyte.
14. The sporogenous tissue is amphithecial in origin.
15. The archesporium develops into fertile spore mother cells and sterile pseudoelaters. The
pseudoelaters may or may not possess spiral thickenings.
16. The capsule starts maturing from the tip downwards.
17. The capsule usually dehisces by two valves.
CLASSIFICATION 8.3
Majority of the bryologists believe that the class Anthoceropsida includes only a single order
(Anthocerotales) having only a single family (Anthocerotaceae). However, Proskauer (1951) and
Reimers (1954) believe that there should be two families (Anthocerotaceae and Notothylaceae) under
the order Anthocerotales.
Only 6 genera and about 300 species are recognised to belong to the class Anthoceropsida. These 6
genera include Anthoceros, Aspiromitus, Dendroceros, Megaceros, Notothylas and Phaeoceros. Four
of these genera (Anthoceros, Dendroceros, Megaceros and Notothylas) are recognised universally
by all the bryologists to belong to Anthoceropsida. Stephani (1916) separated about 55 species of
Anthoceros in the form of a separate independent genus Aspiromitus, while several species of the same
genus (Anthoceros) have been delinked in the form of a new genus, i.e. Phaeoceros by Proskauer
(1951).
ANTHOCEROTACEAE 8.4
1. Plant body is gametophytic and thalloid. The thallus is dark green, dorsiventrally flattened and
lobed. The lobes with divided margins overlap each other.
2. Thallus lacks distinct midrib.
3. The ventral surface lacks scales, tuberculate rhizoids and mucilage hairs.
4. The archegonia are almost completely embedded in the gametophyte.
5. The sporophytes are long, upright, cylindrical structures.
6. At the base of each sporophyte is present a tubular sheath called involucre.
7. The sporogonium is differentiated into a bulbous foot, a meristematic zone and a long
capsule.
8. “Cells of the capsule do not mature at the same rate and the cells in the basal portion of a
capsule remain embryonic even after those in the apical portion are fully mature” (Smith,
1955).
9. A mature capsule dehisces from the apex downwards.
10. Capsule wall is several-layered thick, and its outermost layer contains stomata.
11. Chloroplasts are present in the cells of the sub-epidermal layers of the capsule wall.
12. Columella is present. It is endothecial in origin.
13. Columella remains overarched by archesporium.
14. Simple and branched pseudoelaters are present.
118 � Bryophyta
The life history of Anthoceros is discussed here in some detail.
ANTHOCEROS 8.5
Class—Anthoceropsida
Order—Anthocerotales
Family—Anthocerotaceae
Genus—Anthoceros

Anthoceros is a cosmopolitan genus but occurs mainly in temperate and tropical parts of the world.
Out of a total of about 200 species reported so far from different parts of the world, about 25 species
of Anthoceros have been reported from India. It grows commonly in both plains and hills, on moist
shady places, on the sides of ditches, and also in moist shady hollows among rocks. Dry conditions
are not usually liked by Anthoceros. Some common Indian species, along with their places of common
occurrence, are A. erectus (Kumaon Himalayas, Kulu, Manali, Mussoorie), A. himalayensis (Himalayas
at an elevation of 5000–8000 ft), A. gollani (South India, including Chennai, Travancore), A. longii
(Shimla, Nainital) and A. chambensis (Punjab, Chamba Valley).
GAMETOPHYTIC PHASE
The plant body of Anthoceros, like other bryophytes, is gametophytic, and the gametophytes are
thalloid, dark green, dorsiventral, usually prostrate (Fig. 8.1A-F) and smaller than Marchantia. In a
majority of the species (e.g. A. laevis, A. punctatus), the thalli are variously lobed. The plant body is
somewhat erect or raised over a short stalk in A. erectus, while it is pinnately branched in A. halli, and
somewhat elongated in A. himalayensis.
Margins of the thallus, in most of the species, are irregularly lobed. The branching of the thallus
appears to be of dichotomous type.
The dorsal surface of the thallus may be smooth (A. laevis), or rough with spines or ridges (A.
fusiformis), or velvety because of the presence of several lobed lamellae (A. crispulus).
The ventral surface of the thallus contains only smooth-walled rhizoids. Tuberculate rhizoids and
scales are absent. Mucilage hairs are also not found. Several dark, bluish-green spots are also seen on
the ventral surface of the thallus. These spots are actually the cavities filled with Nostoc, an alga.
8.5.4 Anatomy of Thallus

Anatomically (Fig. 8.2A), the thallus is homogeneous, i.e. shows no differentiation of tissues. Both
the surfaces are marked by an epidermal layer, i.e. upper epidermis on the upper side and the lower
epidermis on the lower side.
Sporogonia
A
Thallus
C
D
Fig. 8.1 A-F, External features of some species of Anthoceros. A, A. punctatus; B, A. laevis;
C, A. erectus; D, A. crispulus; E, A. fusiformis; F, A. himalayensis
In between the two epidermal layers is present a soft, parenchymatous region of more or less uniform
cells. The epidermal cells are comparatively smaller-sized, green, photosynthetic with chloroplasts
somewhat larger than that of the cells of underlying tissues, and are more regularly arranged. The
middle region of the thallus is usually 6 to 8 cells thick in most of the species. But, in a few species, it
becomes 30 to 40 cells thick.
120 � Bryophyta
Fig. 8.2 A, VTS thallus of Anthoceros; B, A mucilage slit on the ventral surface
of the thallus; C, An enlarged cell; D, A much enlarged chloroplast with a pyrenoid.
The air chambers and the air pores are absent in Anthoceros. However, some intercellular mucilage-
filled cavities, each opening externally on the ventral surface by a stomata-like slit (Fig. 8.2B) or slime
pore are present in the ventral region of the thallus. According to Parihar (1961), the slime pores appear
to represent vestigial stomata. Nostoc, a blue-green alga, usually remains inhabited endophytically
in these mucilaginous cavities, and hence these are also called Nostoc - cavities. The mucilage slits
provide entry to the Nostoc filaments inside the thallus. Several schizogenous tubular cavities also
develop in A. punctatus and several other species along with the Nostoc cavities.
Unlike other bryophytes, the cells of Anthoceros are peculiar in that each possesses a single large
chloroplast (Fig. 8.2C), resembling several members of Chlorophyceae (green algae). Some species
have variable number of chloroplasts in their cells. Each cell contains 2 chloroplasts in A. pearsoni
while 4 chloroplasts in A. halli. The chloroplast is also peculiar in possessing a single large pyrenoid,
resembling that of algae (Fig. 8.2D). The pyrenoids of Anthoceros consist of 25 to 300 spindle-shaped
bodies grouped together in the form of a compact body. A rudimentary starch grain is also formed by
each of these spindle bodies of the pyrenoid. The pyrenoids are absent in other members of bryophytes.
The nucleus of each cell is located usually near the pyrenoid in close apposition to the chloroplast.
8.5.5 Apical Growth

The apical growth takes place in the thalli of Anthoceros, but whether it takes place by a single apical
cell or by a group of apical cells is not clear. Mehra and Handoo (1953) opined that in A. erectus and
A. himalayensis the apical growth is initiated by a group of apical cells. On the other hand, Campbell
(1918) observed that the apical growth in A. fusiformis takes place by a single apical cell which cuts off
the segments on dorsal, ventral as well as on both the lateral sides.

Progressive decay and death of older parts of the thallus, formation of tubers, gemmae formation and
survival of only the growing points of the thalli, are the major methods of vegetative reproduction in
Anthoceros.
1. When the progressive death and decay process of the older posterior parts of the thallus
reaches up to the dichotomy, the apical regions of the thallus may survive and start to function
as new plants.
2. Tubers are usually swollen (A. laevis), stalked (A. himalayensis) or unstalked, and storage
perennating bodies which develop during unfavourable conditions, such as prolonged drought.
Other portions of the thallus die during such unfavourable conditions. When moisture and
other favourable conditions are again available, the tuber germinates readily into a new thallus.
Starch grains, aleurone granules and oil globules usually remain present in the inner tissues
of a tuber in Anthoceros. A. halli, A. himalayensis, A. laevis, A. pearsoni and A. tuberosus are
some of the tuber-forming species.
3. Gemmae develop either on the dorsal surface or along the margin of the thallus in several
species, such as A. formosae and A. glandulosus.
4. Except the growing point and a small adjacent portion, the remaining parts of the thallus die
during summer in A. fusiformis. On return of the favourable conditions, the surviving growing
points develop into the new thalli.

The sexual reproduction in Anthoceros is oogamous, as in most of other bryophytes. The species may
be monoecious (e.g. A. fusiformis, A. punctatus) as well as dioecious (e.g. A. dixitianus, A. erectus, A.
halli, A. laevis). The monoecious species are usually protandrous, i.e. antheridia develop first on the
thallus and the archegonia develop later on. The sex organs remain embedded and develop quite close
to the apical cell, i.e. near the growing point.
8.5.8 The Antheridium

1. Development of Antheridium It starts from a superficial dorsal cell which does not become
papillate (Fig. 8.3A). It divides periclinally into an inner antheridial initial and an outer roof initial
(Fig. 8.3B). A mucilage-filled space appears between the roof initial and the antheridial initial. This
space enlarges gradually and forms the antheridial chamber (Fig. 8.3C-G). There is no contribution
of roof initial in the formation of antheridium proper. The roof initial divides anticlinally as well as
periclinally to form a bi-layered roof (Fig. 8.3F-G) outside the antheridial chamber. Further develop-
ment of antheridium differs in different species. In some species (e.g. A. pearsoni), the antheridial
initial develops directly into a single antheridium, whereas in other species it divides by vertical divi-
sions into two, four, eight or more daughter cells, each such cell developing into an antheridium. In
the latter case, several antheridia develop inside an antheridial chamber, as in A. punctatus (Fig. 8.3G).
However, in all the species, further development of antheridium from the antheridial initial is same as
described below:
The antheridial initial first divides by two vertical divisions at right angles to one another, thus
forming four cells. The next division is a transverse division forming an upper tier of four antheridial
cells and a lower tier of four stalk initials (Fig. 8.3D). The stalk initials divide only by transverse
divisions to form a long four-rowed stalk of the antheridium (Fig. 8.3G). Upper four antheridial cells
(Fig. 8.3D) divide transversely to form an eight-celled octant stage (Fig. 8.3E). All the cells of the octant
stage divide periclinally to form eight central primary androgonial cells and eight outer primary
jacket cells. Several rapid transverse and vertical divisions take place in the primary androgonial cells
122 � Bryophyta
Fig. 8.3 A-G, Development of antheridium in Anthoceros punctatus
forming a mass of androgonial cells (Fig. 8.3F,G). All of the androgonial cells ultimately function as
androcyte mother cells. The androgonial mother cells give rise to androcytes. The spermatogenesis
is similar to that of other members of bryophyta. Each of the androcytes metamorphoses into a single
biflagellate antherozoid. The primary jacket cells divide by several anticlinal divisions to form a
single-layered sterile jacket.
Either a single or several antheridia are present in an antheridial chamber. Several antheridia, in
different stages of their development, may be seen developing in an antheridial chamber (Fig. 8.3G).
The secondary antheridia arise as young buds from the base of the mature antheridia. Proskauer (1951)
reported as many as 22 antheridia in an antheridial chamber in A erectus, and up to 25 antheridia in an
antheridial chamber in A. punctatus.
2. Mature Antheridium It is a stalked, club-shaped (Fig 8.4A) or pouch-like body, surrounded

by a single-layered sterile jacket. The stalk usually consists of four rows of cells. Each cell of the jacket
contains a well-developed plastid. The sterile jacket encloses several androcytes, each of which meta-
morphoses into a biflagellate antherozoid.
Fig. 8.4 A, A mature antheridium of Anthoceros laevis; B-C, Antherozoids of A. laevis;

D, An antheridium of A. punctatus showing dehiscence
3. Antherozoids Each antherozoid is a unicellular, uninucleate and biflagellate structure having a

linear body (Fig. 8.4B, C) with a slightly broader head. The length of the flagella is almost the same as
that of the body of the antherozoid. An elongated swelling, probably representing a blepharoplast, is
present just beneath the flagella.
4. Dehiscence of Antheridium The mature antheridia soon become exposed due to the irregu-
lar breaking of the roof of the antheridial chamber. The antheridia now start absorbing water. Soon,
some of the cells of the apical region are separated from each other forming an apical aperture in the
antheridium. The mass of the androcytes come out through this apical aperture (Fig. 8.4D).
8.5.9 The Archegonium

1. Development of Archegonium The archegonia are produced in acropetal succession near
the growing points. The archegonia remain embedded in the thallus. Each archegonium is characteristi-
cally surrounded externally by a mucilaginous mound (Fig. 8.5).
The development of archegonium starts from a superficial dorsal cell which starts to function as
an archegonial initial (Fig. 8.6A). Differing from Marchantia, the archegonial initial does not project
from the surface of the thallus. Mehra and Handoo (1953) reported that the archegonial initial starts to
function directly as the primary archegonial cell. (Anthoceros thus differs from Marchantia because
in the latter, the archegonial initial first divides transversely into an outer primary archegonial cell and
an inner stalk cell).
124 � Bryophyta
Fig. 8.5 An embedded archegonium of Anthoceros laevis surrounded by a mucilaginous mound
Fig. 8.6 A-G, Archegonial development in Anthoceros (B2 is the T.S. of B1)
In Anthoceros, the primary archegonial cell divides by three vertical intersecting divisions, forming
three peripheral initials or jacket initials and a central primary axial cell (Fig. 8.6B1, B2). The
primary axial cell divides transversely into two equal-sized cells called outer cell and primary ventral
cell (Fig. 8.6C). The primary ventral cell soon becomes distended, whereas the outer cell divides
transversely into an upper cover initial and an inner primary neck canal cell (Fig. 8.6D). The cover
initial divides by one or two vertical divisions at right angles to one another to form two or four cover
cells (Fig. 8.6E). The primary neck canal cell divides by transverse divisions to form four to six or
more neck canal cells (Fig. 8.6E, F). The primary ventral cell divides by a transverse division to form
a ventral canal cell and an egg (Fig. 8.6E).
Along with all these developments, the three jacket initials divide transversely to form two tiers of
cells (Fig. 8.6C). The three cells of the upper tier, surrounding the neck, divide by vertical divisions to
form six vertical rows of cells. These vertical rows of cells surround the neck canal cells. Owing to the
embedded nature of the archegonium, further development of the lower tier of the cells of the jacket
initials is indistinguishable from that of the surrounding cells of the thallus.
2. Mature Archegonium Except that of the cover cells, the major parts of the mature archego-
nium remain embedded in the tissues of the thallus. The axial row consists of four to six neck canal
cells, a venter canal cell and an egg (Fig. 8.6F). Soon, the cover cells are thrown off without leaving any
of their traces. The neck canal cells and venter canal cell disintegrate and form a mucilaginous mass,
leaving only the egg in the cavity of the venter of the archegonium (Fig. 8.6G).
3. Fer liza on It is exactly similar to that of Marchantia discussed under Article 7.7.9.
SPOROPHYTIC PHASE
8.5.10 Development of Sporogonium
Immediately after fertilization, the fertilized egg enlarges in size and keeps on enlarging till it completely
fills the cavity of the venter. A cellulose wall is secreted outside this developing zygote (Fig. 8.7A).
Usually, the zygote first divides by a vertical division (Fig. 8.7B) to form two equal cells. It is followed
by a transverse division in each cell forming four equal- or unequal-sized cells (Fig. 8.7C). If the four
unequal-sized cells are formed after first two divisions, then the two basal cells are smaller while
the two upper cells are larger (Fig. 8.7C). In the four-celled embryo, the next division is a vertical
division at right angles to the first vertical division. Thus, an eight-celled octant stage of the embryo
is resulted. According to Bhardwaj (1950), however, the first division in the zygote is a transverse
division, followed by two successive vertical divisions at right angles to one another, thus forming an
eight-celled octant stage.
Further development of the eight-celled embryo is different in different species. In Anthoceros
erectus, the lower four cells develop into foot whereas the upper four cells form the intermediate
zone (seta) and capsule. In A. himalayensis and A. pearsoni, however, the upper four cells of the eight-
celled embryo divide by transverse walls to form a three-tier embryo (Fig. 8.7D) in which each tier
contains four cells. Of these three tiers, the uppermost tier forms the capsule, the middle tier forms the
intermediate zone (seta) and a small part of the foot, while the lowermost tier of four cells forms the
remaining major part of the foot.
Foot develops further by regular (A. erectus) or irregular (A. himalayensis) divisions in the cells of
the lowermost tier of the embryo. It ultimately becomes broad, bulbous and multicellular (Fig. 8.7G,
H). Its cells are parenchymatous. In most of the species, the superficial cells of the foot tend to become
rhizoidal or haustorial. These cells penetrate deeply into the adjoining tissues of the thallus and absorb
food, and thus behave like cells of haustorium.
126 � Bryophyta
The capsule develops from the uppermost tier of four cells. This uppermost tier divides by one or
two transverse walls to form the cells arranged in two or three tiers containing four cells each (Fig.
8.7E). These cells now divide periclinally to form an outer layer of amphithecium and an inner layer
of endothecium (Fig. 8.7E). The entire endothecium, consisting of four vertical rows of cells, gives
rise to columella. The columella in the mature capsule consists of 16 vertical rows of cells. In A.
fusiformis, the columella in the mature sporogonia becomes very massive and consists of as many as
36-49 rows of cells, according to Campbell (1924).
Fig. 8.7 A-H, Development of the sporogonium in Anthoceros; I-M, LS of the mature capsule through
different por ons of the sporogonium (Note the single-layered archesporium in I, bilayered
archesporium in J, forma on of spore mother cells and elater mother cells in K, forma on
of spore tetrads in L, and forma on of spores and pseudoelaters in M)
A periclinal division in the amphithecium divides it into an outer sterile layer of the initials of jacket
layer and an inner fertile layer of the sporogenous tissue called archesporium (Fig. 8.7G). The jacket
layer initials divide periclinally several times to form a four- to six-layered wall of the capsule (Fig.
8.7H-M). The outermost layer of the wall of the capsule functions as epidermis. In mature capsule,
the epidermis is thickly cutinised and also bears some pore-like stomata. The cells of the other wall
layers are green, parenchymatous and bear intercellular spaces. The number of chloroplasts in the
parenchymatous cells of the inner wall layers may vary from one to four in different species.
The archesporium overarches the apex of the columella in young sporogonia (Fig. 8.7H). Several
stages of the archesporium development may be seen and studied in a single sporogonium. At the
base of the capsule, the archesporium is single-layered and present in between the columella and the
wall layers (Fig. 8.7I). In species, such as A. himalayensis and A. pearsoni, the archesporium becomes
bilayered slightly above the base (Fig. 8.7 J). In A. hallii, it even becomes 2 to 4 cells in thickness.
At a slightly higher level in the maturing sporogonium, the archesporium gets differentiated into two
types of cells, i.e. fertile spore mother cells and sterile pseudoelater mother cells (Fig. 8.7K). The
spore mother cells are oval to spherical, large cells with dense granular cytoplasm, chloroplast and
prominent nucleus. The pseudoelater mother cells are elliptical cells with less prominent nuclei. In
several species, the spore mother cells and pseudoelater mother cells are alternately arranged. Both are
soon separated from each other as well as from the columella due to the unequal growth of the wall
layers and the columella.
The spore mother cells are diploid in nature. Each enlarges in size, becomes globular and divides
reductionally to form four haploid spores, arranged in a tetrad manner (Fig. 8.7L). The pseudoelater
mother cells divide either transversely or by oblique divisions to form a loose net of sterile cells, which
are soon separated into one- to three-celled pseudoelaters (Fig. 8.7 L, M; Fig. 8.8). When young, the
pseudoelaters contain starch and protein, and are nutritive in function. But mature pseudoelaters are
dead cells and help in the dispersal of spores.
The middle tier of the young embryo (Fig. 8.7D) usually gives rise to the intercalary meristem,
which functions like that of the seta of Marchantia and other Hepaticopsida. After the formation of the
columella, the archesporium and the jacket, further growth of the sporogonium takes place mainly due
to the activity of this intercalary meristem (Fig. 8.7H, I). Due to activity of this meristematic region,
the capsule increases in length, its tip portion is dehisced, and ultimately the mature spores are liberated
(Fig. 8.7M).
The calyptra or involucre protects the young developing sporogonium in the form of a sheath or
fleshy covering. It is formed mainly by the thallus tissues surrounding the young sporogonium and
partly by the tissues of the embedded archegonium. When the sporogonium grows more, the involucre
is ruptured and ultimately remains only in the form of a sheath at the base of the sporogonium (Fig.
8.7I). Structurally, the involucre resembles the thallus because it is mainly the part of the latter.

It consists of a bulbous foot, an intermediate meristematic zone in place of seta, and an erect cylindrical
capsule, varying between 2 and 15 centimetres in length in different species (Fig. 8.7I-M). The foot is a
bulbous body consisting mainly of parenchymatous cells and some superficial cells showing haustorial
processes. The latter penetrate deep into the tissues of the thallus and absorb food.
128 � Bryophyta
The capsule is a long structure, having sterile columella and fertile sporogenous tissue or
archesporium. The columella extends from the base up to the tip of the capsule and consists of 16
vertical rows of cells. Campbell (1924) opined that the columella functions as a water-conducting
tissue of the capsule in A. fusiformis. However, it mainly provides mechanical support to the long
sporogonium and helps in spore dispersal.
The columella is surrounded by sporogenous tissue. It consists of a one-layered archesporium at
the base, while mature spores and pseudoelaters are at the top of the capsule. The pseudoelaters are
multicellular and branched or unbranched structures. Spiral thickenings are absent in pseudoelaters
(Fig. 8.8). The capsule wall consists of 4 to 6 layers of cells, of which the outermost layer is the
epidermis. The continuity of the epidermis is broken by a few pore-like stomata.
A B
Fig. 8.8 A-B, Pseudoelaters of Anthoceros erectus (A) and A. himalayensis (B)
8.5.12 Dehiscence of the Capsule

In a mature capsule, the tip becomes brownish or blackish in colour and ultimately shrinks due to the
loss of water in the dry conditions. Because of this shrinkage, the capsule wall comes in contact with
the fertile sporogenous region containing spores and pseudoelaters. Actual dehiscence occurs because
of slits or dehiscence lines formed in different species. Spiral twisting is also observed in the tip region
of the capsules because of the excessive loss of water. Enlarging columella and somewhat hygroscopic
nature of the pseudoelaters also help in the dehiscence of the capsule. Spores are dispersed over a long
distance from the dehiscing capsule by strong air currents.
GAMETOPHYTE
8.5.13 Spore
The haploid spores, formed after the reduction division of the diploid spore mother cells (Fig. 8.9 A-B)
are the starting points of the gametophytic generation. Each spore is surrounded by a thin endospore
and a thick outer layer of exospore. The spores may have short papillae on the outer surface or may be
reticulate with furcate spines. Each spore is uninucleate and usually contains oil globules, a plastid and
other food materials. The diameter of the spore in different species usually ranges between 0.03–0.06
mm. The colour of the mature spores may be yellow, dark brown or smoky black.

Spore germination has been studied only in a few species of Anthoceros, such as A. fusiformis
(Campbell, 1928), A. erectus and A. punctatus (Mehra and Kachroo, 1962). The exospore layer of
the spore ruptures along the triradiate ridge (Fig. 8.9A) and the endospore comes out in the form of
a germ tube (Fig. 8.9C). The plastid, oil globules and other food materials of the spore pass into the
Fig. 8.9 A-H, Spore germina on in Anthoceros
germ tube. Usually, the first two divisions in the germ tube are transverse, forming a three-celled short
filament (Fig. 8.9D, E). The uppermost cell now divides by a vertical division (Fig. 8.9F). Soon, the
same type of vertical division takes place in the lower cell (Fig. 8.9G). All the so-formed four cells
now divide by yet another vertical division at right angles to the first one, forming an eight-celled stage.
Further growth of the young gametophyte (Fig. 8.9H) usually takes place by the four upper cells of this
eight-celled germling. A cylindrical elongated young gametophyte is soon resulted. The first rhizoid
develops from the young multicellular gametophyte, from any cell containing chloroplast.
8.5.15 Life History of Anthoceros

Life history of Anthoceros is depicted in Fig. 8.10.
APOSPORY IN ANTHOCEROS 8.6

“Production of a diploid gametophyte from vegetative cells of the sporophyte, that is, without the
production of spores is known as apospory. It was first reported in Anthoceros laevis by Lang (1901).
He discovered that pieces cut off from the sporogonium of this species, when placed in conditions
suitable for their growth, are able to develop into masses of cells which grow into thalli. Commonly,
such masses of cells develop from sub-epidermal cells. Since such a gametophyte was produced directly
from the vegetative cells of the sporophyte, this clearly represents a case of the phenomenon of apospory.
Later on, apospory was reported in several species of Anthoceros. Schwarzenbach (1926) observed that
in many species of this genus, any cell of the young sporophyte can produce a gametophyte, except
that of the cells of the meristematic region. The gametophytes producted aposporously appear quite
normal to that of other normal thalli, except the fact that the number of chromosomes in their cells is
diploid (2x).
130 � Bryophyta
S
Y G
N F
G I
C Zygote Two-celled
Egg D A
M embryo
Egg inside mature Y H
Archegonium
archegonium
SPOROPHYTE
2N
PARASITIC
J
E
Antherozoid
B Young sporogonium
GAMETOPHYTE Columella
Antheridium N
INDEPENDENT
M
A
E K
Mature thallus
I
O
S Portion of
LS of sporogonium
I
S L
O Spore
M mother cell (2X)
Young thallus N Spore
Spore tetrad
(X)
Fig. 8.10 Life history of Anthoceros
NOTOTHYLACEAE 8.7
1.The sporophytes grow out horizontally from the fertile lobes of thallus.
2.In comparison to Anthocerotaceae, the sporophytes in Notothylaceae are shorter, more
compact and marginal in position.
3. Capsule wall lacks photosynthetic tissue.
4. The outermost layer of capsule wall lacks stomata.
5. In some species of this unigeneric family, columella is absent. In others, however, it is well-
developed.
6. Pseudoelaters are simple, and are equal-sized or sometimes larger than the spores.
7. Pseudoelaters contain spiral or oblique bands.
The only genus of the family Notothylaceae is Notothylas, and some details of its life history are
discussed here.
NOTOTHYLAS 8.8
Systema c Posi on
Class—Anthoceropsida
Order—Anthocerotales
Family—Notothylaceae
Genus—Notothylas
About a dozen species of Notothylas have been reported so far, growing in damp, shady places,
either on moist soil or on rocks or on moist floor walls of old buildings. They are widely distributed
in temperate as well as tropical regions. Five species have been reported so far from India. These are
Notothylas chaudhurii, N. indica, N. javanicus, N. levieri and N. pandei. Two most common Indian
species, growing in Himalayas, are N. indica and N. levieri. N. indica also occurs commonly in the
plains.
Thallus Sporogonium Upper epidermis
Antheridia
Sporogonium
Thallus
Rhizoids
B Lower epidermis
C
VS of thallus
A Thallus bearing sporophytes
of Notothylas indica
Thallus bearing sporophytes
of Notothylas levieri
Capsule wall
Involucre layers
Sporogonium
Columella
Capsule Spores
Foot Columella
Thallus Elaters Foot
Meristem
D
E F
Foot G
Thallus showing enlarged
sporogonium Thallus showing dehisced
LS of sporogonium LS of sporogonium sporogonium
of Notothylas indica of Notothylas levieri
Fig. 8.11 A-G, Notothylas. A, Thallus of N. levieri bearing sporophytes; B, Thallus of N. indica bearing
sporophytes; C, Ver cal sec on of the thallus; D, Thallus showing an enlarged sporogonium; E, LS
sporogonium of N. indica; F, LS sporogonium of N. levieri; G, Thallus showing dehisced sporogonium
132 � Bryophyta
Plant body is gametophytic, prostrate, thalloid, delicate, light green and orbicular or sub-orbicular
in outline. The thalli are variously lobed, and the lobes are either toothed, seriate or fimbriate (Fig.
8.11A-B). Thalli of Notothylas indica form bluish-green rosettes while that of N. levieri are pale green,
elongated and irregularly branched. N. indica thalli attain a diameter of 2–5 mm. Horizontally borne
sporophytes and cyanobacterial auricles are present on the dorsal surface of the thallus. Only smooth-
walled rhizoids are present on the ventral surface. Tuberculate rhizoids and scales are absent.
Anatomy of the thallus reveals that it is 6–8 cells thick in the middle and only one or two cells thick
on the margins (Fig. 8.11C). Cells of the upper and lower epidermal layers are generally smaller than the
other inner cells. Soft parenchymatous cells are present in between the two epidermal layers. Mucilage
cavities containing the Nostoc colonies are common in the thallus. These colonies are, however, absent
in Notothylas javanicus.
The structure and development of sex organs in Notothylas (Fig. 8.12 A, B) is exactly similar to
Anthoceros, as described earlier under Articles 8.5.7 to 8.5.9.
Fig. 8.12 Notothylas. A, Antheridia in different stages of development; B, An archegonium bearing young
embryo; C, Three- ered embryo; D-E, Differen a on of outer and inner amphithecial layers and
endothecium.
The zygote first divides by transverse division in N. indica and N. orbicularis. In some other species
(N. javanicus and N. levieri), however, the first division in the zygote is a longitudinal division. In all
species of Notothylas, further divisions result in the development of a three-tiered embryo, in which
each tier contains four cells (Fig. 8.12C). The lower two tiers develop into the foot, while the uppermost
tier of this three-tiered embryo produces the capsule. Periclinical division in the cells of the uppermost
tier demarcates the outer amphithecium and inner zone of endothecium. The embryo development is
similar in all species up to this stage only. After this stage, it is different in different species.
In Notothylas indica, the amphithecium cells divide periclinally and form outer amphithecium
and inner amphithecium. The outer amphithecium forms the wall of the capsule, while the inner
amphithecium forms the sporogenous tissue. The endothecium develops into columella (Fig. 8.12
D-E). In this way, N. indica resembles Anthoceros in its embryo development. In N. levieri, however,
the entire amphithecium develops into the wall of the capsule while the entire endothecium develops
into sporogenous tissue. Thus, the embryo development in N. levieri resembles with members of
Hepaticopsida. This species, therefore, serves as a connecting link between Anthoceropsida and
Hepaticopsida. In N. indica, therefore, the archesporium is a layer of cells in between the capsule wall
and columella. The archesporium becomes many-layered towards the apex of the sporophyte in this
species.
Regarding the fate of archesporium in Notothylas, two bands of sterile and fertile tissues are formed.
Bands of the sterile tissue form elaters and of fertile tissue form spore mother cells. Structurally,
the elaters are short, often curved, unicellular structures with bands of thickenings on their walls.
The diploid spore mother cells divide reductionally and form haploid spores. A meristematic zone
of intercalary meristem is present at the base of the capsule. The meristematic zone is short-lived in
activity and contributes to various tissues in the capsule. Due to this short-lived activity of meristematic
tissue, the sporophyte in Notothylas is shorter (only 2–3 mm in length) than that of Anthoceros.
Mature sporophytes (Fig. 8.11 D-F) of Notothylas are pointed, oval or cylindrical structures. Usually,
they are cylindrical with pointed ends. When young, the sporophyte is completely enclosed within a
membranous structure. Each sporophyte has a massive foot, meristematic zone and capsule. The foot is
triangular and some of its basal cells elongate to form rhizoidal outgrowths. Intermediate meristematic
zone is short, less active and exhibits very little meristematic growth. Capsule remains surrounded by
a 3–4 layered wall, which lacks stomata and photosynthetic tissue. Columella is present in the centre.
The sporogenous tissue contains elaters and haploid spores. The spores are dark brown in colour, and
their wall is differentiated into thick exospore and thin endospore. One to four lines of dehiscence in
the form of rows of thick-walled cells can be observed. They are seen running along the entire length
of the capsule, and they meet at the apex of the capsule.
Dehiscence of capsule and dispersal of spores take place along lines of dehiscence. At the time of
spore germination, a four or eight-celled mass of cells is formed. It is called germ disc. Rhizoids start
developing from one or two basal cells of the germ disc. Differentiation of a marginal meristem is
responsible for the further growth of the thallus of Notothylas.
BIOLOGICAL IMPORTANCE OF THE SPOROPHYTE

OF ANTHOCEROS 8.9
The sporophyte of Anthoceros is remarkably different from the sporophytes of other bryophytes, and
it is thus considered unique and also of advanced type. Its advanced characteristics show evolutionary
trends.
8.9.1 Some Advanced Features of Sporophyte

1. Capsule Wall of the Sporophyte The wall of the capsule of the sporophyte of Anthoceros
is multilayered, contains chlorophyll-containing green cells with several intercellular spaces between
134 � Bryophyta
them. On the external side is present a layer of epidermis with several stomata, here and there, like
that of higher plants. The presence of amply ventilated photosynthetic tissue in the wall is an advanced
feature, and certainly a step towards the beginning of physiological independence of the sporophyte.
To some extent, it can prepare its own food. Inspite of this, it never becomes completely independent
of the gametophytic thallus.
2. Decentraliza on and Steriliza on of the Central Fer le Region, i.e.

Endothecium Differing from many other bryophytes, the archesporium is amphithecial in origin
in Anthoceros. Decentralization and complete sterilization of the central fertile tract is very clear in
this genus. The central core of sterile tissue forms the columella in the capsule. The columella is made
up of narrow, vertically elongated conducting cells, the walls of which are uniformly thickened. The
presence of central columella in the sporophyte of Anthoceros suggests of the origin of the vascular
tissue in plants. Vascular elements (xylem, phloem, etc.), however, never develop in columella. But its
central location in the sporophyte may be compared with the vascular cells of the early tracheophytes.
Like vascular tissue of higher plants, the columella in Anthoceros provides mechanical support to the
sporophyte. Some scientists believe that presence of columella is an evolutionary step towards the
development of protostele.
Archesporium, being amphithecial in origin, is also an evolutionary steps because the spore-
producing region is shifted from central to the superficial region, and this position promotes easy
dispersal of spores.
3. Importance of Development of Archesporium into Alternate Bands of Fer le and

Sterile Tissues Fertile spore mother cells and sterile pseudoelater mother cells develop in between
columella and wall layers in alternate bands in the capsule of Anthoceros. This is also an evolutionary
tendency, which has great potentialities. It is thought by some bryologists to be an initial step towards
the origin of sporangia and sporophylls. Bower thought it to be a step towards the origin and evolution of
leaves and sporangia in pteridophytes. This theory of Bower is called theory of origin of strobilus.
4. Presence of Basal Intercalary Meristem Presence of intercalary meristem at the capsule

base provides strength and also unlimited power to the entire sporophyte to grow. The meristem keeps
on adding new cells at the base of the capsule. These cells keep on differentiating into columella,
archesporial region and also photosynthetic region of capsule wall. All these activities of meristematic
cells prolong the period of spore production. Only because of these characteristics, Anthoceros sporo-
phyte is long-lived in comparison to other bryophytes, and keeps on producing spores as long as the
gametophytic thallus is surviving.
5. Massive Foot Foot is well-developed, massive and remains embedded in the thallus. It pro-
duces many short, rhizoid-like processes, which penetrate into the thallus and keep on absorbing the
nutrients. Presence of rhizoid-like processes in the foot is an advanced feature which also indicates
evolutionary trends.
6. Upright Cylindrical Body of the Capsule The upright cylindrical body of the capsule
helps in the easy dispersal of spores, and it is also thus an advanced feature of the sporophyte in
Anthoceros.
8.9.2 Anthoceros Sporophyte and Origin of Pteridophytes

The above-mentioned six unique and advanced features of the sporophyte of Anthoceros also indicate
the evolutionary trends or possible lines of biological progress in this genus. On the basis of such
characteristics, Campbell (1940) formulated the well-known theoery of Anthocerotalean Origin of
Pteridophytes. In this theory, “the sporophyte of Anthoceros and its allies was considered to be on the
line of evolution leading to the simplest and primitive independent sporophyte of the pteridophytes”
(Campbell, 1940). In Anthoceros fusiformis Campbell also collected some very large and bulky
specimens of the sporophyte, attaining a length of about 15 cm or even more, and suggested that these
have evolved into some simplest and most primitive free-living sporophytes of pteridophytes. Such
large sporophytes of A. fusiformis also survived in unusually favourable habitats for a large period
of as long as 9–10 months or even more. The following peculiarities were seen in such specimens of
A. fusiformis:
1. The sporogenous tissue was suppressed or ill-developed at the base of the capsule of the
sporophyte.
2. In comparison to normal sporophytes, there was seen amply ventilated photosynthetic system
in the capsule wall of A. fusiformis.
3. Bulky growth of the columella was seen. It became nearly double in diameter in A. fusiformis
than that of the normal sporophytes of other species of Anthoceros.
4. Some columella cells at the base of the capsule became elongated, functioned like conducting
strands and thus can be compared with the simple vascular bundles of tracheophytes.
5. Bulky and unusually large foot may come in direct contact of the soil due to disorganisation
of the adjacent tissue of the thallus.
These characteristics of A. fusiformis suggest that such sporophytes have attained a “condition
comparable to that of the young pteridophyte after it has established its first root” (Campbell, 1940).
This theory of Anthocerotalean Origin of Pteridophytes has, however, been criticised by several other
later bryologists.
AFFINITIES OF ANTHOCEROTALES 8.10

Available details of life history (gametophytic and sporophytic generations) of Anthocerotales, in
general, and Anthoceros, in particular, lead us to the conclusion that this group has characteristic
features common with algae on one hand and other groups (like Hepaticopsida, Bryopsida and also
pteridophyta) on the other. Let us have a glimpse of some features of Anthocerotales common with all
these other groups.
8.10.1 Features Common with Algae

Anthocerotales show features common with algae, in general, and green algae (Chlorophyceae), in
particular. Some of such features are listed below:
1. Presence of at least a single large chloroplast in each cell of the gametophyte. In Megaceros,
however, many chloroplasts are present in each cell, and this has been interpreted as an
intermediate condition between the single chloroplast of Anthoceros and many chloroplasts of
other Embryophyta.
136 � Bryophyta
2. Presence of pyrenoid in the chloroplast of each cell (Pyrenoid is actually a small grain of
protein in the chloroplast of an algal cell, around which starch is deposited. Pyrenoids are the
characteristics of cells of green algae, i.e. Chlorophyceae).
3. Presence of simple, green, thallus-like gametophytic plant body, resembling in outline and
branching.
4. Presence of biciliate antherozoids in both algae and Anthocerotales.
All these above-mentioned similarities suggest that Anthoceros is close to the evolutionary line
leading from algae (Chlorophyceae) to the land plants.
8.10.2 Features Common with Hepa copsida

Anthocerotales show many features common with Hepaticopsida, yet another major group of
bryophytes. Some of such features are listed below:
1. Presence of simple, thallus-like gametophyte in both Anthocerotales and most of the species
of Hepaticopsida.
2. Appendages are absent on the thalli of both.
3. In both Anthocerotales and many Hepaticopsida, there is not much differentiation of tissues in
the plant body.
4. Apical growth is almost similar in members of both groups.
5. There exists similarity in the essential construction of mature sex organs in members of both
the groups.
6. Both Anthocerotales and Hepaticopsida possess biflagellate antherozoids.
7. In both groups, amphithecium and endothecium separate in almost the same way by periclinal
divisions.
8. In both the groups, archesporium gives rise to fertile spores and sterile elaters or
pseudoelaters.
9. In Megaceros, walls of the sterile cells of the capsule are spirally thickned to form elaters,
quite similar to that of members of Hepaticopsida.
10. Notothylas levieri is a noncolumellate species of Anthocerotales. The entire endothecium in
this species forms the archesporium, and the amphithecium forms the jacket of the capsule.
Due to these similarities, N. levieri forms a connecting link between Anthocerotales and
Hepaticopisida.
8.10.3 Features Common with Bryopsida or Mosses

Features of sporophyte of Anthoceros common with that of mosses (Bryopsida) are listed below:
1. Highly differentiated and ventilated photosynthetic region is present in the capsule wall of
both Anthoceros and mosses.
2. Columella, the central solid core of sterile cells, is present in both Anthoceros and Funaria.
3. Greatly reduced sporogenous tissue present only in a small part of the capsule in both.
4. Presence of functional stomata in the wall of both Anthoceros and Funaria.
5. Anthoceros resembles some Bryopsida (e.g. Sphagnum) in their common characteristic of
origin of archesporium from inner amphithecium. In some species of Notothylas, however,
the archesporium is endothecial in origin, and this characteristic may form a link between
Bryopsida and Anthocerotopsida.
8.10.4 Features Common with Pteridophyta

Some characteristics of Anthocerotales common with pteridophytes include the following:
1. Overall general similarity in the thalli (gametophytes) of Anthocerotales and prothallus of ferns.
2. Deeply sunken sex organs in the thalli of Anthocerotales and prothallus of several pteridophytes.
3. Similarity in the structure of mature female sex organs, i.e. archegonia.
4. Similarity in the sporophyte of Anthocerotales with the rootless and leafless dichotomously
branched sporophytes of some fossil pteridophytes belonging to Psilophytales.
8.10.5 Anthocerotales: A Synthe c Group

The characteristics outlined above under Articles 8.7.1 to 8.7.4 suggest that Anthocerotales are a distinct
but synthetic group of plants. It forms a bridge or connecting link between the (i) Hepaticopsida and
Bryopsida (mosses) on one hand, and (ii) bryophytes and pteridophytes on the other. Simultaneously,
Anthocerotales also have some features common with green algae (i.e. Chlorophyceae). In this
connection, SK pande (1932, 1934), a noted Indian bryologist, suggested that Notothylas levieri
forms a connecting link between Anthocerotales and Hepaticopsida. Pande, however, opined that both
Anthocerotae and Hepaticae should be retained as two separate independent classes of Bryophyta.
DIFFERENCE BETWEEN ANTHOCEROTOPSIDA

AND HEPATICOPSIDA 8.11
Some striking differences between Anthocerotopsida and Hepaticopsida are listed below in Table 8.1:
Table 8.1 Difference between Anthocerotopsida and Hepa copsida
S. NO. ANTHOCEROTOPSIDA HEPATICOPSIDA

1. Each cell contains a single chloroplast. Numerous chloroplasts are present in each cell.
2. Each chloroplast contains a pyrenoid. Pyrenoids are absent in the chloroplast.
3. Sex organs are deeply sunken in the thallus. Sex organs are superficial in nature, and not deeply
sunken.
4. A very narrow meristematic zone is present in the Meristematic zone is absent in sporogonium (e.g.
sporogonium in between foot and capsule. Riccia), or it is replaced by a well-differentiated
seta in between foot and capsule (e.g. Marchantia).
5. Columella is present as a slender core of sterile Columella is absent.
tissue in the capsule.
6. Archesporium is originated from amphithecium. Archesporium is endothecial in origin.
7. An amply ventilated photosynthetic system is Amply ventilated photosynthetic system is absent
present in the capsule wall of the sporogonium. in the capsule wall of the sporogonium.
8. Stomata are present in the outermost layer of capsule. Stomata are absent in the capsule wall.
9. Pseudoelaters, without spiral thickenings, are Instead of pseudoelaters, the elaters with spiral
present. thickenings are present (Marchantia).
10. The usual chromosome number is 5 or 6. 10. The number of chromosomes are usually 8 or 9.
138 � Bryophyta
A NOTE ON THE ORIGIN OF ANTHOCEROTALES 8.12

Origin of Anthocerotae is still difficult to be finalised because of the fundamental differences in the
genetic relationship of this group and Hepaticopsida. Campbell (1925) believed that the nearest affinities
in the thallus and sporophyte of Anthocerotae are with some pteridophytes. The view of Campbell
was supported by Kashyap (1929) and proved by showing several striking structural similarities
between the thallus of Anthoceros erectus and prothallus of Equisetum debile. Kashyap (1929)
tried to prove that Anthocerotae originated from Equisetum debile by reduction. In 1953, PN Mehra
and ON Handoo opposed the views of Campbell and Kashyap and gave several reasons to support
their contention. Some of these reasons include the following: (i) There are basic and fundamental
differences between the sporophytes of Anthocerotae and pteridophytes, (ii) Vascular tissues (xylem
and phloem) are completely absent in Anthocerotae while present in all pteridophytes, and (iii) Vast
difference in the basic chromosome number of Anthocerotae and pteridophytes, 5 in Anthocerotae
but 10 in Equisetum debile. They, however, proposed that Anthocerotae and Rhyniaceae have sprung
up from the same ancestral stock, i.e. Anthorhyniaceae in pre-Devonian (Mehra and Handoo, 1953).
The Anthorhyniaceae, according to them, evolved from a hypothetical pro-liverwort stock from which
have also arisen Marchantiales, Jungermanniales and Sphaerocarpales. Pro-liverwort stock originated
in some Chlorophyceae (green algae). GM Smith (1955) has shown the phylogenetic relationship
between Anthocerotae and Psilophytales (e.g. Rhynia and Horneophyton) of pteridophytes, which are
some of the oldest and most primitive of all the known vascular plants. Further evidence in favour
of Psilophytalean ancestry of Anthocerotales was given by J Proskauer (1962) on the basis of his
studies of Dendroceros crispus (of Anthocerotales) and Rhynia and Horneophyton (of Psilophytales).
Professor Ram Udar and DK Singh (1978) supported the views of Proskauer on the basis of their
studies on thickened bands in the capsule wall of Notothylas levieri. Their research has been published
in The Bryologist.
1. What are Anthocerotes?

2. Who was the first to propose the name Anthocerotopsida to Anthoceropsida?
3. Anthocerotopsida members are commonly named as _____.
4. Write at least seven general characteris cs of Anthoceropsida.
5. Whether Anthoceropsida members contain tuberculate rhizoids?
6. Are scales in Anthoceropsida present or absent?
7. Each cell of Anthoceropsida contains at least one chloroplast and one _____.
8. Instead of seta, the sporophyte of Anthocerotes contain _____?
9. In Anthoceropsida, the sporogenous ssue is _____ in origin.
10. Two families of Anthocerotales are Anthocerotaceae and _____.
11. Six genera of Anthoceropsida are Anthoceros, Aspiromitus, Dendroceros, Phaeoceros,
Notothylas and _____.
12. Write a brief note on the classifica on of Anthoceropsida.
13. Write at least five characteris c features of Anthocerotaceae.
14. Write a short note on the distribu on and habitat of Anthoceros with par cular reference to
India.
15. Write an illustrated essay on the life history of Anthoceros in about 1000 words.
16. Thalli of which of the following are smaller: Marchan a or Anthoceros?
17. Give an illustrated account of the anatomy of thallus of Anthoceros in about 200 words.
18. Which of the following are present in thallus of Anthoceros?
(a) Stomata (b) Scales (c) Nostoc colonies (d) Tuberculate rhizoids
19. Explain means of vegeta ve reproduc on in Anthoceros.
20. Describe development of antheridium in Anthoceros making suitable diagrams.
21. What do you mean by protandrous?
22. In Anthoceros, the archegonia remain embedded in the thallus or projected on the thallus?
23. Give a detailed accout of development of sporogonium in Anthoceros.
24. Explain structure of a mature sporogonium of Anthoceros.
25. Write a short note on the pseudoelaters of Anthoceros.
26. Depict life history of Anthoceros with suitable illustra ons only.
27. Write a note on the apospory in Anthoceros.
28. Write any four characteris cs of the family Notothylaceae.
29. Write a detailed scien fic note on the life history of Notothylas in about 500 words.
30. Discuss the biological importance of the sporophyte of Anthoceros.
31. Write some advanced features of the sporophyte of Anthoceros and explain their
implica ons.
32. Give a detailed account of the affini es of Anthocerotales.
33. Write some such features of Anthocerotales which are common with Hepa copsida and
pteridophytes.
34. Anthocerotales is a synthe c group. Explain
35. Write some of the basic differences between Anthocerotopsida and Hepa copsida.
36. Write a note on the origin of Anthocerotales.
9
Bryopsida
(General Account)
WHAT IS BRYOPSIDA? 9.1
Bryopsida or Musci is a class of bryophytes that includes mosses. It includes different evolutionary
lines, most prominent amongst which is Bryales, and due to this it has been given the name Bryopsida.
The class Bryopsida includes about 660 genera and 15000 species. These are the bryophytes which
have a leafy (not thalloid) gametophyte with the leaves not strictly in 2 or 3 ranks, multicellular rhizoids
and, in most, a capsule (sporophyte) with both a columella and a lid (operculum).
The gametophyte in Bryopsida is the dominant generation and exhibits two distinct morphological
stages. The first, which arises on germination of the spore, is the filamentous protonema, which,
except for its oblique cross walls, resembles a heterotrichous green alga. The protonema produces
buds, from which the familiar leafy moss plant arises.
Although the group does not generally have a very rich fossil record but earliest fossil mosses are
seen in the rocks of Permian.

1. Bryopsida or Musci includes the bryophytes called “mosses”.
2. Mature gametophyte is foliose, i.e. divisible into rhizoids, stem and leaves. It is never thalloid.
The gametophyte may be acrocarpous or pleurocarpous. In acrocarpous gametophytes,
the main axis is terminated by the development of the reproductive organs, and so subsequent
growth is sympodial. In such mosses, the main axis is almost always erect. On the other hand,
in pleurocarpous gametophytes, the reproductive organs are produced laterally and the main
axis is usually creeping.
3. The rhizoids are multicellular, well-branched, and contain oblique septa or cross walls.
4. The leaves are spirally arranged on the stem.
5. The sex organs are present in the apical portions of the gametophore.
6. The sporogonium consists of foot, seta and spherical or a cylindrical capsule.
Bryopsida (General Account) � 141
7. The sporogonium is determinate in growth.

8. The seta elongates gradually.
9. The capsule wall is several-layered, and its outermost layer is the epidermis containing some
functional or nonfunctional stomata.
10. The capsule opens by operculum or by four slits.
11. A well-developed sterile columella is usually present in the centre of the capsule.
12. The lower part of the capsule may be green and photosynthetic.
13. The sporogenous tissue or archesporium develops either from the endothecium or from the
amphithecium, and usually encloses the columella.
14. Only spores develop from the archesporium. The elaters or other sterile cells are usually
absent.
15. The capsule dehisces in dry weather, and the dehiscence is often controlled by hygroscopic
peristomial teeth.
16. The spore germinates usually into an extensive and filamentous protonema.
CLASSIFICATION 9.3
Bower(1935) and Campbell (1940) divided the class Bryopsida into three orders, viz. Bryales,
Sphagnales and Andreaeales.
Bryales are commonly called true mosses. It is the largest order of about 600 genera including
Polytrichum, which shows some internal differentiation. Sphagnales are commonly termed bog or
peat mosses. It contains the single genus, Sphagnum, characteristic of waterlogged acid areas. The
third order is Andreaeales, which also contain only one genus Andreaea, the members of which are
known as granite mosses.
Engler et al. (1954) divided Bryopsida into five subclasses, i.e. Sphagnidae, Andreaeidae, Bryidae,
Buxbaumiidae and Polytrichidae.
However, Parihar (1965) and Holmes (1986) divided the class Bryopsida (Musci or mosses) into
three subclasses, viz. Sphagnidae (peat or bog mosses), Andreaeidae (granite mosses) and Bryidae
(true mosses).
For details of the classification proposed by Holmes, refer Chapter 2, Article 2.1. For latest views
on classification of mosses, consult details proposed by Goffinet and Buck (2004), and Troitsky et al.
(2007) as outlined in Chapter 2, Article 2.3.2.
DISTRIBUTION AND HABITAT 9.4

Mosses are distributed throughout the world and in almost all types of habitats, except in seas and oceans.
A few species of mosses have even been reported from very high altitudes, as much as 20000 feet above
the sea level (e.g. Aongstroemia julacea). Mosses, however, flourish most in the wet and humid regions,
and prefer to grow in moist plains and mountain forests of tropical and subtropical regions. Sphagnum
and a few more mosses grow in bogs. (A bog is a region of badly drained permanently wet land that is
subject to high rainfall and has a persistently moist atmosphere). A few mosses are even aquatic (e.g.
Fontinalis antipyretica). Some have been reported even from deserts (e.g. Torula desertorum). Majority
of mosses, however, grow in damp situations and form extensive mats.
142 � Bryophyta
HABIT 9.5
Plant body of mosses is gametophytic, and gametophytes are green and independent. Mosses can be
divided into the following two broad categories on the basis of their habit:
(a) Acrocarpous mosses, in which the main axis is almost always erect.
(b) Pleurocarpous mosses, in which the main axis is usually crreping.
In acrocarpous mosses, the main axis terminates by the development of the reproductive organs,
and thus the subsequent growth is sympodial. But in pleurocarpous mosses, the main axis does not
terminate by the reproductive organs, which are therefore produced laterally.
9.5.1 Growth Phases of Moss Gametophyte

Protonema stage and leafy stage (Fig. 9.1A-D) are the two growth phases of the moss gametophyte.
1. Protonema Stage This stage of mosses is creeping, green, branched, multicellular and often
filamentous. It develops from the spore. It is a vegetative phase and bears no sex organs. It is also called
juvenile stage. In a majority of mosses, the protonema dies and disappears soon, thus making the leafy
gametophytic plants independent. In some mosses, however, the protonema persists for a long time
(e.g. Buxbaumia aphylla).
2. Leafy Stage It is the adult stage of moss plant, and due to the presence of leaves it is called
leafy stage. The gametophyte in this stage consists of an upright, slender axis, bearing spirally arranged
leaves. Sex organs develop on the leafy stage. Actually, it is the so-called moss plant, which we see
in the form of gametophyte. Leafy stage develops as a lateral bud from the protonema stage. From the
single protonema stage of the mosses develop many leafy moss plants.
Fig. 9.1 Some stages of the development of protonema and leafy stages. A, A young germina ng
spore; B, Protomena; C, Protonema and ini als of gametophore of Splachnum ovatum;
D, Protonema with mature male and female gametophores of Ephemerum serratum.
9.5.2 Moss Gametophore

Moss gametophore remains differentiated into a stem-like axis bearing many green, leaflike expansions.
Some prefer to call them stem and leaves while others term cauloid to the stem and phylloids to the
leaves. The plant body remains attached to the substratum by many brown-coloured filaments known as
rhizoids. Three major or basic organs of a moss gametophore are stem, leaves and rhizoids.
The stem or central axis is branched or unbranched, and erect or prostrate in different species. The
branching, if present, is usually lateral and never dichotomous.
The leaves are green, sessile and the main photosynthetic organs of the gametophore. The basal
part of the leaves is usually broad. The form of the leaf is highly variable in different species. It may
be ovate to linear or sub-orbicular. The cells of the leaves remain filled with chloroplasts. Each leaf
contains a midrib. In some mosses, however, the midrib is absent (e.g. Sphagnum). The lamina of each
leaf is usually one cell in thickness. Basically, the leaves remain arranged spirally in three ranks on the
stem, and, therefore, the gametophore has radial symmetry. Typical 3-ranked spiral arrangement with
1/3 divergence is seen in Fontinalis, but this arrangement may be different in different genera, e.g. 2/5
in Sphagnum, 3/8 in Funaria and 5/13 in Polytrichum.
The rhizoids are multicellular, well-branched and septate. The septa in moss rhizoids are oblique.
They keep the plants attached to the substratum, and their main function is anchoring. Rhizoids are
brown or dark brown in colour. They develop in tufts in pleurocarpous mosses.
VEGETATIVE OR ASEXUAL REPRODUCTION 9.6

Mosses have exceptional ability to reproduce vegetatively, and that is how they form dense mats over
large areas. Almost all parts of moss gametophore (rhizoid, axis, leaf) or any portion of all of these parts
have the capacity to reproduce vegetatively. Some such methods of vegetative or asexual reproduction
in mosses include (i) branching of leafy stems, (ii) formation of stolons, (iii) detachment of a specially
modified branch of bud-like form as in Pohlia, (iv) formation of lateral buds from the extensively
branched primary protonema, (v) secondary protonema, (vi) persistent apices, (vii) gemmae, which
are green, oval, multicellular buds produced on short stalks (e.g. species of Torula, Leptobryum and
Barbula), and (viii) tubers, which are underground resting buds (e.g. Funaria, Bryum etc.).
APOSPORY IN MOSSES 9.7

Production of a diploid gametophyte from vegetative cells of the sporophyte, that is, without the
production of spores is called apospory. Several mosses show extraordinary power of regeneration
through apospory. Any undamaged cell of the moss sporophyte, besides the gametophyte, can grow
into a protonema-like body. Green protonema-like filaments are commonly produced by wounding of
the unspecialised cells of any of the parts of sporophyte. Many leafy gametophytes are developed from
such green protonema-like filaments. Thus, the moss plants are produced directly from the sporophyte
without the production of spores under the phenomenon of apospory. Such plants have a diploid
chromosome number, instead of haploid. They are quite normal like that of haploid plants, except that
they are slightly larger-sized. They also produce gametes, but such gametes are not haploid but diploid.
Their fusion, under the process of fertilization, results in the formation of a tetraploid sporophyte.
144 � Bryophyta
However, if such a diploid gamete fuses with a normal haploid gamete, it results in the formation of a
sporophyte which is triploid (i.e. possesses three sets of chromosomes).
SEX ORGANS 9.8

9.8.1 Distribu on of Sex Organs
Two sex organs (antheridia and archegonia) are borne in clusters, generally at the top of the main axis or
lateral branches. However, in some (e.g. Sphagnum) the antheridia occur singly in the axil of perigonial
leaves. Several sterile green filaments (paraphyses) are also present intermixed with sex organs in
each cluster. Mosses are monoecious as well as dioecious. Monoecious mosses (e.g. Mnium medium)
possess male and female sex organs on the same individuals. But dioecious mosses (e.g. Buxbaumia)
possess male and female sex organs on different individuals of the same plant species.
9.8.2 Categories of Monoecious Mosses

The following three different categories can be recognised in monoecious mosses on the basis of the
distribution of sex organs:
1. Paroicous Mosses These are the monoecious mosses in which two different sex organs de-
velop in the same cluster or head in separate groups. A few perichaetial leaves separate groups of two
sex organs.
2. Autoecous Mosses These are the monoecious mosses in which two different sex organs de-
velop on separate branches of the same plant.
3. Synoicous Mosses These are the monoecious mosses in which two different sex organs de-
velop in the same cluster or head intermingled with each other.
Some mosses show four conditions, viz. paroicous, autoicous, synoicous and dioecious (e.g. Pohlia).
9.8.3 Antheridia
Moss antheridia are elongate, club-shaped structures with a short and multicellular stalk (Fig. 9.2 A).
They are comparatively longer and narrower than that of antheridia of liverworts. Their size ranges
between 0.2 mm and 2.00 mm, with an average length of 1.5 mm. Mature antheridia are usually orange
in colour and remain surrounded by a jacket layer enclosing a mass of androcytes or male gametes. The
tip of each antheridium usually contains one to many large-sized cells. Each male gamete or sperm is a
spirally coiled, motile, biflagellate, uninucleate and unicellular structure.
9.8.4 Archegonia
Except that of a comparatively longer stalk, massive venter and longer neck, the archegonia of mosses
(Fig. 9.2B) are similar to those of liverworts. Each archegonium consists of a long neck and a globular
venter. The neck encloses 35 to 50 neck canal cells, a ventral canal cell and an egg. The primary cover
cells are present at the tip of the archegonium.
Fig. 9.2 Sex organs of mosses. A, An antheridium of Funaria hygrometrica;

B, An archegonium of Cyathophorum bulbosum
9.8.5 Fer liza on

Fertilization is effected through the agency of water. In paroicous and synoicous mosses, the presence
of a connecting film of water between archegonia and antheridia is a necessity. In autoecious and
heterothallic mosses, the antherozoids are transported up to the archegonia generally through water.
It has been finally established that antherozoids enter the archegonium in response to the chemical
stimuli. Union of male and female gametes (i.e. plasmogamy) is followed by karyogamy (i.e. fusion
of two nuclei), and thus a diploid zygote is resulted.
9.8.6 Embryo and Sporophyte

A wall is soon secreted around the fertilized egg. The so-formed zygote increases in size, divides and
redivides to form an embryo. The embryo is actually the product of repeated mitotic divisions of the
zygote. The growth of the moss embryo is due to the activity of two apical cells located in opposite
directions for quite some time. The apical cell, situated at the upper end of the embryo, is more active
and from its derivatives are differentiated the capsule and a large part of the seta. The foot and lower
remaining portion of the seta are differentiated from the derivatives of the lower apical cell. The young
embryo is long and slender. The lower end of the embryo burrows through the stalk of the archegonium
and into the apex of the gametophore. Simultaneously, there is an enlargement of the surrounding
archegonium, which is known as calyptra.
The sporophyte of most of the mosses consists of foot, seta and capsule. It resembles those of
Jungermanniales in showing the formation of a long seta, which elevates the capsule above the
surrounding leaves of the gametophore. In most mosses, the seta contains a well-defined central strand.
Its main function is conduction of food material to the developing capsule. Seta also functions as a
mechanical tissue. The foot is the basal part of the sporophyte. Its functions include attachment and also
as an absorbing organ. The foot remains embedded in the tissues of the tip of the leafy gametophore.
Capsule is the uppermost, most complex region of the sporophyte of mosses. In true mosses (e.g.
Funaria hygrometrica), it is divisible into three major regions, viz. apophysis, theca and operculum
(Fig. 9.3). Apophysis is the basal region, theca is the middle region and operculum is the uppermost
region of the capsule.
146 � Bryophyta
Fig. 9.3 Longitudinal sec on of the capsule of Funaria hygrometrica
Apophysis is the green photosynthetic region, containing several chlorenchymatous cells and
stomata in its outermost layer. Theca is the spore-producting region of the capsule. It consists of
several parts including outermost epidermis, wall layers, air cavities traversed by trabeculae, spore
sacs and centrally-located columella. The archesporium is endothecial in origin. Sporogenous tissue
forms spore mother cells, which divide meiotically to form haploid spores. Elaters are absent. The
uppermost region of capsule matures into operculum and peristome. The peristome contains teeth-
like projections that surround the mouth of the capsule. The number of peristomial teeth varies between
4 and 64, but in various mosses they are in multiples of 4, such as 8, 16, 32 or 64.
In some mosses (e.g. Polytrichum), the peristomial teeth are solid structures made up of bundles of
dead cells. Such a peristome is known as nematodontous peristome. They are arranged in a single
series and they do not show any hygroscopic movements. In some other mosses, the peristome is made
up of thin, membranous teeth made up of thickened portions of cell walls of the adjacent cells. Such
a peristome is known as orthodontous peristome, and their teeth are hygroscopic in nature. They are
either arranged in single series or in two series. When arranged in one series, the peristome is known
as haplolepidous, but if arranged in two series, the peristome is known as diplolepidous (e.g. Funaria
hygrometrica).
9.8.7 Spore and its Germina on

Moss spore is a unicellular, uninucleate structure surrounded by a thin spore wall. The reserve food
is in the form of lipid and a little starch. Mitochondria and endoplasmic reticulum and rarely Golgi
bodies are also present in the cytoplasm of the spore. Three wall layers, which constitute the spore wall
are perine, exine and intine from outside within. A fourth separating opaque layer may or may not be
present. When present, it is actually a sub-unit of the exine.
Spores start germinating immediately if they fall on a suitable soil. In some species, however, the
length of time varies if the conditions are unfavourable. Acording to Meyer (1941), in some species,
the spores remain “capable of germinating eight to sixteen years after shedding”. At the time of
germination, a spore increases somewhat in size, ruptures the outer spore wall layer and then sends out
one or two germ tubes. A cross wall is soon formed near the point of emergence of the germ tube. The
cell thus cut off by this cross wall soon develops into a branched, multicellular, filamentous structure
known as protonema. Two types of branches are soon differentiated in this protonema. These are (i)
chloronemal branches or chloronema, which usually grow along the surface of the substratum or into
the air for some time, and (ii) rhizoidal branches or rhizoids, which penetrate into the substratum. The
chloronemal branches have (i) colourless cell walls, (ii) septa at right angles to lateral walls, and (iii)
several well-defined chloroplasts. On the other hand, rhizoidal branches possess (i) brown cell walls,
(ii) oblique or diagonal septa, and (iii) ill-defined chloroplasts or leucoplasts. A number of the leafy
gametophores develop as lateral outgrowth from the protonema. They bear sex organs and represent
the adult stage of moss gametophyte.
Most of the genera of mosses have a disappearance of protonema after leafy gametophores produced
by them have attained a stage where they have rhizoids and several leaves. In a few genera, however,
protonema persists throughout the entire life of the gametophytic generation. Protonema functions as
the major photosynthetic portion of the gametophyte in some mosses, e.g. Ephemerum (Fig. 9.1D).
RESEMBLANCES AND DIFFERENCES BETWEEN HEPATICOPSIDA

LIVERWORTS AND BRYOPSIDA MOSSES 9.9
9.9.1 Resemblances
There are only a few resemblances between liverworts and mosses:
1. Both show heteromorphic type of alternation of generations.
2. Both lack meristematic tissue in their sporogonia. Some meristematic tissue, however,
develops in Anthocerotopsida.
3. Plant body is gametophytic in both.
4. Both possess biflagellate antherozoids.
5. Members of both groups possess calyptra.
9.9.2 Differences between Hepa copsida and Bryopsida

Differences between the two groups are many. Some such differences are tabulated in Table 9.1.
148 � Bryophyta
Table 9.1 Differences between Hepa copsida and Bryopsida
S. NO HEPATICOPSIDA (LIVERWORTS) BRYOPSIDA (MOSSES)

1. Development of gametophytic plant body from a Development of gametophytic plant body from a
spore is a continuous process. spore is not a continuous process. It passes through
two phases, viz. (i) juvenile stage which includes
formation of protonema from spore, and (ii) adult
leafy gametophore stage.
2. Spore first germinates into a sporeling, which Spore first develops into a well-branched fila-
develops into a mature gametophytic plant body. mentous protonema, from which develop many
gametophores.
3. Plant body is dorsiventral and thalloid or foliose. Plant body is radial and always foliose.
4. Plant body of most liverworts lacks a central Central conducting strand is present in the axis of
conducting strand. mosses.
5. Leaves, present in foliose liverworts, lack midrib. Leaves of all mosses have a midrib.
6. Growth of leaves in foliose members is usually Growth of leaves in mosses is by the activity of an
intercalary. apical cell.
7. Scales are present on the ventral surface in thal- Scales are absent in mosses.
loid liverworts.
8. Rhizoids are unbranched and unicellular. Rhizoids are well-branched and multicellular.
9. Oblique septa are absent in the rhizoids. Rhizoids of most mosses have oblique septa.
10. Sex organ development is intercalary and not Sex organ development is due to the activity of an
apical. apical cell.
11. Antheridia are ovoid or subglobose in shape. Antheridia are longer, narrower and club-shaped bodies.
12. Archegonial neck contains lesser number (usu- Archegonial neck is very long and contains more
ally 6–9) neck canal cells. (usually 35–45) neck canal cells.
13. Early growth of the embryo is intercalary. Early growth of embryo is usually biapical.
14. Seta, when present, is soft and lacks any internal Seta is long, tough and contains a central conduct-
differentiation of tissues. ing strand.
15. Seta lengthens, and due to this the sporophyte Seta lengthens, and due to this the sporophyte
breaks through the calyptra, but quite late. breaks through the calyptra at an early stage.
16. Sporophyte is quite simple in its organization. Sporophyte is quite complex in its organisation.
17. Sporophyte lacks apophysis and operculum In most mosses, the sporophyte contains regions
region. like apophysis and operculum.
18. Stomata are absent on the capsule wall. Capsule wall of mosses contains stomata.
19. Sporophyte lacks annulus. Capsule of most mosses contain annulus.
20. Capsule lacks columella. Columella is present in moss capsule.
21. Sporophyte lacks peristomial teeth. Peristomial teeth are present.
22. Elaters are present. Elatere are absent.
23. Usually, the entire endothecium is archesporial Usually, only the outermost layer of endothecium is
in origin. archesporial in origin.
1. What is Bryopsida? Explain in about 100 words.

2. Bryopsida includes bryophytes which are commonly called _____.
3. Write at least ten general characteris cs of Bryopsida.
4. Do Bryopsida have thalloid gametophores?
5. In mosses, a spore germinates into a mul cellular, branched, filamentous structure known as
_____.
6. In mosses, the gametophyte may be acrocarpous or _____.
7. What are acrocarpous gametophytes in mosses?
8. In the centre of the moss capsule, a well-developed sterile region is present. It is known as
_____.
9. Elaters in mosses are _____.
10. Write a short scien fic note on the classifica on of Bryopsida.
11. Sphagnales are commonly known as peat or _____ _____.
12. The name “granite mosses” is given to the members of _____.
13. What are true mosses?
14. Write a note on the distribu on and habitat of mosses in about 100 words.
15. Do mosses occur in seas or oceans?
16. What are the two growth phases of moss gametophyte? Explain briefly.
17. Name five major types of the vegeta ve reproduc on found in mosses.
18. Write a brief scien fic note on apospory in mosses.
19. How will you differen ate between the following?
(a) Paroicous mosses, (b) Autoecous mosses, (c) Synoicous mosses
20. Draw labelled diagrams of an antheridium and an archegonium of any moss.
21. What is calyptra?
22. Differen ate between apophysis, theca and operculum of a moss capsule.
23. What is peristome?
24. How can you differen ate between nematodonous peristome and orthodontous
peristome?
25. Describe structure and germina on of moss spore in about 200 words.
26. Enumerate any four resemblances between liverworts and mosses.
27. What are the ten major differences between Hepa copsida and Bryopsida? Tabulate them.
10
Bryopsida
(Selected Mosses)
SPHAGNALES OR BOG MOSSES 10.1
10.1.1 What are Bog Mosses, Bogs and Peat?
Sphagnales are commonly known as “bog mosses” or “peat mosses”. They are characteristic of
waterlogged acid areas.
Bog is actually a region of badly drained permanently wet land that is subject to high rainfall and
has a persistently moist atmosphere. Bogs are commonly found in upland and waste areas of temperate
regions. Besides several angiospermic plants like rushes (e.g. Juncus) and sedges (e.g. Carex), the most
common bryophyte of bogs is Sphagnum.
Peat is actually the partially decomposed plant material. It builds up in areas with poor drainage,
namely bogs and fens. Acid bog peat or “peat moss” is composed primarily of the remains of bog plants
such as Sphagnum mosses and sedges.
10.1.2 Differences between Bog Moss and Other Mosses

Bog moss differs from other mosses in possessing the following:
1. Thalloid protonema which develops into gametophore
2. No midrib in the leaves
3. Leaves usually composed of markedly different types of cells
4. Axillary antheridia, which show a clear differentiation of fertile part
5. Acrogynously formed archegonia
6. Archesporium is amphithecial in origin
7. Pseudopodium in the sporophyte, which is an elongation of a stalk of gametophytic tissue.
Bryopsida (Selected Mosses) � 151
SPHAGNUM 10.2
10.2.1 Distribu on, Habitat and Some Other Details
Sphagnum is the only genus of Sphagnales, which contains only one family Sphagnaceae. It is commonly
known as bog moss. Some bryologists also name Sphagnum as turf moss or peat moss.
Sphagnum is a cosmopolitan genus which grows from north and south tropics through the temperate
regions and extends up to sub-arctic and sub-antarctic regions. It is particularly abundant in the northern
circumpolar regions. It occurs as dense masses in ponds, lakes and other such surroundings where due
to seepage such soft water is available which contains only a little amount of lime. In cooler temperate
regions of the northern hemisphere, Sphagnum usually dominates the vegetation of wetlands. Over 350
species of Sphagnum have been reported so far, of which were than 20 species occur in India. The pH
of the water in which Sphagnum grows ranges from 3.7 to 4.9. The size of Sphagnum plants varies from
a few inches to a maximum of 7 inches.
Sphagnum is a perennial moss and keeps on growing year after year. Water around Sphagnum plants
is so much acidic that there develops only a little decay of dead basal portions. Regularly, the older
parts of the plants of Sphagnum die and the dead organic remains of these plants, in combination with
the remains of other surrounding plants, form a compact mass known as peat. Due to the formation
of peat, this moss is known as peat moss, and being
a peat-former, it is of great commercial importance
for us.
10.2.2 Mature Gametophore

Structurally, the plant is erect, branched and
differentiated into stem and leaves. The rhizoids are
colourless and develop at the base but do not survive for
too long. They disappear soon. Mature gametophores
usually lack rhizoids. At the apex of the gametophore,
there are present a number of short branches densely
crowded in a cluster, called coma (Fig. 10.1A). In the
posterior part of the stem, the branches arise in tufts
in the axil of every fourth leaf, and in each tuft, there
are 3 to 8 branches. The branches are of two types
(Fig. 10.1A, B): (i) divergent branches, which are
short, stout and grow outward and upward, and (ii)
drooping branches, which hang downward quite
close and around the stem. These are also called
flagelliform branches. In submerged plants, these
drooping branches are absent. At certain intervals,
one of the branches in the tuft grows and develops
into an apical cluster of branches like the main stem.
This is known as innovation. On being separated Fig. 10.1 Sphagnum. A, A part of gametophore;
B, Por on of a plant showing tu of
from the main branch, the innovation develops into a branches
152 � Bryophyta
new plant, and hence it functions as an organ for vegetative propagation. The first-formed leaves are in
three vertical rows or 3-ranked. The arrangement, however, changes to 2/5 in the later stages. The stem
is only a few inches in length, and close aggregation of short and stumpy branches towards the apex
(Fig. 10.1 A) provides it an appearance like that of a head or capitulum. An exceptional feature of the
Sphagnum leaves is the absence of a midrib.

Terminal growth of the stem is due to a
tetrahedral apical cell with three cutting faces
(Fig. 10.2). Each segment is first cut off by a
periclinal division giving rise to an inner cell
and an outer cell. The inner cell forms the axis
of the shoot while the outer cell gives rise to
the cortex and a single leaf. In this fashion,
each segment develops into a single leaf
and the subtending part of the stem. When
young, the leaves are in three vertical rows,
corresponding to the three cutting faces of
the tetrahedral apical cell. But during course
of time, as the stem and leaves grow, the
Fig. 10.2 Longitudinal sec on through the apex of a
leaves show some displacement, and they gametophore of Sphagnum subsecundum.
finally loose their 3-ranked arrangement. At
maturity, the leaves get arranged on the stem
in a spiral with a divergence of 2/5, i.e. each leaf gets separated from the next leaf above it by 2/5th of
the circumference of the stem.
10.2.4 Leaves
Leaves show an orderly arrangement of two types of
cells (Fig. 10.3A), viz. (i) green, narrow and elongate
chlorophyll-containing cells, and (ii) hyaline or colourless,
large, polygonal cells which lack cell contents. In the cross
section of a mature leaf (Fig. 10.3B), the large and dead
hyaline cells, and green, wedge-shaped chlorophyllous
cells are present alternately. Thickening bands and pores
are present in hyaline cells. With reference to size, shape
and structure, the leaves of the main stem are different
from those present on the branches. The leaves present on
the main axis or stem have very little or no chlorophyll,
and their hyaline cells lack thickening bands and pores. On
the other hand, the leaves present on the branches consist
of a network of green, long, chlorophyll-containing cells. Fig. 10.3 Sphagnum. A, A part of mature leaf
Some of these green cells surround one hyaline dead showing green and hyaline cells;
porose cell. B, Sec on of a leaf showing green
and hyaline cells
10.2.5 Anatomy of Axis

Transverse sections of the axis (Fig. 10.4 A-D) show an outermost layer of cortex made up of compactly
arranged cells. In some species, the cortex becomes 3-5 cells thick. Mature cortical cells get devoid of
protoplast, become hyaline and dead. In some other species (e.g. S. cymbifolium), they develop spiral
thickenings and even develop pores. Such cortical cells contain water and air. In S. molluscum and S.
tenellum, some cortical cells become greatly enlarged and flask-shaped. They accumulate water and are
known as retort cells (Fig. 10.4B, C). Such type of cortex is known as hyalodermis. The cells inner
to the cortex are prosenchymatous and provide mechanical support to the stem. This region of axis is
known as hadrome (Fig. 10.4A). In such stems, the innermost region is known as medulla. Medulla
cells are vertically elongated, colourless and parenchymatous.
Fig. 10.4 Sphagnum. A, TS Young stem; B, Part of stem showing retort cells; C, TS stem of S.
molluscum showing retort cells; D, TS of old stem showing thickening band and conduc ng
zone
154 � Bryophyta
10.2.6 Reproduc on
Sphagnum reproduces vegetatively and sexually.
1. Vegeta ve Reproduc on
Special branches, known as innovations, are the means of vegetative reproduction. They develop in the
axis of leaves on the main axis in a similar fashion like ordinary branches. They are strong and remain
directed towards the upper side. They get separated because of the progressive death of the basal part
of the main axis and establish themselves as the new plants of Sphagnum.
Some species of Sphagnum multiply by primary protonema. Few marginal cells of the thalloid
primary protonema become meristematic and develop into a multicellular filament. The apical part of
this multicellular filament develops into a thallus-like, flat secondary protonema. A leafy gametophore
of Sphagnum may develop from any of the marginal cells of this secondary protonema.
2. Sexual Reproduc on
Sphagnum has both monoecious as well as dioecious species. The archegonia and antheridia develop
on different branches of the same plant in monoecious species. Development of sex organs takes place
usually in the beginning of the winter season. They develop on specialised branches, formed in the axis
of leaves. Sex-organ-bearing branches are comparatively much smaller than that of the other vegetative
branches of the plant
(a) Antheridium
(i) Antheridial or Male Branches The male branches are catkin-like, small structures (Fig. 10.5 A),
arranged usually spirally or in straight rows on the main axis. They possess many small, coloured
leaves. The colour of these leaves may vary from yellow or brown or even reddish. In the axil of each
leaf of the antheridial branch develops an antheridium (Fig. 10.5B). The development of antheridia in
the antheridial branch is in acropetal succession, i.e. oldest antheridium is present at the base and the
youngest at the apex of the antheridial branch.
(ii) Development of Antheridium It starts from a superficial cell of the antheridial branch. This is
known as antheridial initial (Fig. 10.6A), which soon enlarges and appears like a papillate outgrowth.
The antheridial initial first divides by some transverse divisions to form a small filamentous structure
(Fig. 10.6B). Its terminal cell divides by two intersecting divisions, and thus differentiates an apical
cell (Fig. 10.6B) with two cutting faces. It forms its derivatives in both the planes, resulting into a 10-
to 15-celled structure. Upper 2–5 cells of this structure differentiate into the body of the antheridium
while the remaining lower cells form the stalk of the antheridium. The upper cells, responsible for the
formation of the antheridium body divide periclinally to form outer jacket initials and inner primary
androgonial cells (Fig. 10.6C). The jacket initials divide and form a single-layered jacket. The primary
androgonial cells divide and redivide in different planes to form many androgonial cells. The last
generation of these cells forms androcytes (Fig. 10.6C-G). The antheridium of Sphagnum is unique in
its characteristic that its apical cell stops dividing eventually and starts functioning as a cell of jacket.
Each androcyte metamorphoses into a unicellular, spirally coiled, biflagellate antherozoid.
Fig. 10.5 Sex organs of Sphagnum. A, A lateral branch bearing antheridial and archegonial
branches; B, Longitudinal sec on of antheridial branch
(iii) Mature Antheridium and its Dehiscence The mature antheridium (Fig. 10.6G) has a long stalk
and a globular body. The stalk is made up of 2-4 row of cells. The body is surrounded by a single
layer of sterile jacket, which encloses a mass of androcyte cells. Each androcyte metamorphoses into a
biflagellate, unicellular, uninucleate antherozoid (Fig. 10.6H).
The mature antheridium dehisces and liberates the antherozoids. Apical cells of the sterile jacket
absorb water and become swollen. Due to this, the turgor pressure increases and the wall of the
antheridium breaks into several irregular lobes or valves at the apex. These valves turn backward, and
thus the mass of androcytes is exposed and finally liberated. During this process, the antherozoids come
out of the androcytes and start swimming in the surrounding liquid.
(b) Archegonium
(i) Archegonial or Female Branches Archegonial branches (Fig. 10.5A) are shorter than antheridial
branches. They are bud-like structures bearing either a single or a group of 2–5 archegonia surrounded
by leaves. These leaves are green and larger than those of the leaves of vegetative branches. The leaves
which surround the archegonia are known as perichaetium (Fig. 10.7).
Two types of archegonia develop in an archegonial branch. The archegonium, which develops from
the apical cell of the archegonial branch, is situated at the top and known as primary archegonium.
On the other hand, the archegonia, which develop from the derivatives of the apical cell, are known
as secondary archegonia (Fig. 10.7). However, the process of the development of both the types of
archegonia is the same.
156 � Bryophyta
Fig. 10.6 A-H Sphagnum showing development of antheridium
(ii) Development of Archegonium The development of

archegonium starts from an archegonial initial, which
divides by repeated transverse divisions to form a short
filament of 4–6 cells (Fig. 10.8A-C). The uppermost or
terminal cell of this filament divides by three intersecting
vertical divisions, resulting in the formation of three
peripheral jacket initials and a central primary axial
cell (Fig. 10.8D). The primary axial cell now divides
transversely to form an upper cover initial and a lower
central cell (Fig. 10.8E). Vertical divisions in the cover
cell result into a group of eight or more cover cells, which
form the jacket of the upper or terminal part of the neck
of the archegonium (Fig. 10.8F,G, J). Simultaneously,
the lower central cell also divides transversely to form
Fig. 10.7 LS of apical por on of female branch
an upper primary neck canal cell and a lower primary
bearing archegonia and perichae um
venter cell (Fig. 10.8F). Repeated transverse divisions take place in the primary neck canal cell and
thus develops a row of 8–10 neck canal cells (Fig. 10.8 J). Side by side, the primary venter cell divides
by only one transverse division to form an upper venter canal cell and a lower egg (Fig. 10.8J). Side
by side, each of the three jacket initials (Fig. 10.8 H) divides anticlically as well as periclinally to form
the middle and basal parts of the jacket of the archegonium (Fig. 10.8J). Now, jacket cells divide by
repeated transverse divisions to ultimately form five or six vertical rows of neck cells (Fig. 10.8 H,J).
The jacket becomes two- to three-layered in the basal parts of the neck due to periclinal divisions.
(iii) Mature Archegonium The mature archegonium of Sphagnum (Fig. 10.8J) has a long stalk, a
twisted neck and a massive venter. The neck jacket is two- to three-celled thick in its basal and middle
parts. Eight or sometimes more cover cells are present in the apical part of the neck. The neck cavity
contains 8–10 neck canal cells, and the venter contains a venter canal cell and an egg.
Fig. 10.8A-J Development of archegonium in Sphagnum

158 � Bryophyta
(c) Fer liza on Fertilization takes place only in the presence of water. The liberated antherozoids keep
freely swimming in this water and reach near the neck of the archegonium. Inside the archegonium,
the neck canal cells and venter canal cell disintegrate and disorganise, and form a passage for the
antherozoids. Only the egg is now present inside the venter. One of the antherozoids fuses with this egg
and the fusion product results in the formation of a diploid egg.
(d) Sporophyte
(i) Development of Sporophyte The diploid zygote (Fig. 10.9A) is the first cell of the sporophytic
generation. A single sporophyte usually develops on an archegonial branch. The zygote enlarges and
soon divides by a transverse wall to form a lower hypobasal cell and an upper epibasal cell (Fig.
10.9B). Its further fate is different in different species. In Sphagnum subsecundatum, both the epibasal
as well as hypobasal cells divide by repeated transverse divisions to form a young filamentous embryo
of 6–12 cells (Fig. 10.9C). In some other species (e.g. S. acutifolium), only the epibasal cell of the
bicelled embryo divides transversely, while the hypobasal cell divides by a vertical division, followed
by many irregular divisions resulting ultimately into the formation of a bulbous parenchymatous foot
(Fig. 10.9E-I).
In the filamentous embryo (Fig. 10.9C), usually the 3 or 4 upper cells give rise to the capsule,
and from the remaining cells develop the foot and seta (Fig. 10.9C-E). The upper cells divide by two
vertical divisions at right angles to each other, and thus a quadrant is resulted from each cell, They
divide periclinally to form the outer or peripheral amphithecium and inner or central endothecium
(Fig. 10.9J). Repeated transverse divisions of the endothecium give rise to columella, which forms
the central sterile part of the capsule. The amphithecium divides periclinally to form an outer sterile
layer, and an inner fertile layer of archesporium. The archesporium is thus amphithecial in origin
(Fig. 10.9I-L). A 3–7-layered capsule wall is resulted as a result of periclinal divisions of the outer
sterile layer. The archesporium forms a dome-shaped arch over the columella (Fig. 10.9I). It divides
periclinally to form a 2–4-layered sporogenous tissue, of which all cells start functioning as spore
mother cells. They divide meiotically to form haploid spores, which remain enclosed in a spore sac
in the capsule.
(ii) Mature Sporophyte The mature sporophyte contains foot, seta and capsule (Fig. 10.10A-B). The
foot is made up of parenchymatous cells. It is a bulbous or cylindrical body. Foot in Sphagnum is
haustorial in function. The seta is ill-developed, inconspicuous and has a very narrow structure. The
capsule is well-developed, quite conspicuous and has a spherical structure (Fig. 10.10B). The mature
capsule is a dark-brown or black-coloured body, surrounded by a 2–7 layered wall, of which the
outermost layer is differentiated into an epidermis bearing several nonfunctional stomata. Several
cells of the capsule wall contain chloroplasts, which shows photosynthetic nature of the capsule. At the
apical part of the capsule is present the operculum. It is a circular, biconvex disc-shaped structure. A
circular groove of thin-walled cells separates the operculum from the capsule. It is known as annulus.
In mature sporophytes, the operculum gets separated from the annulus and due to this the spores are
dispersed
Fig. 10.9 A -Development of sporophyte (Note that J-L are the stages of differen a on of
amphithecium and endothecium in transverse sec ons)
Columella is the central part of the capsule. It is made up of sterile cells. A dome-shaped arch of
fertile sporogenous tissue is present over the columella. The calyptra and perichaetium surround the
sporophyte in young conditions.
At maturity, the axis of the archegonial branch elongates, and the capsule thus comes out of the
protective covering of calyptra and perichaetium. The leafless, elongated axis of the archegonial
branch, present at the base of the sporophyte, is known as pseudopodium (Fig. 10.10A). It is mainly a
post-fertilization development. A sac-like structure, formed by the distal end of the pseudopodium and
basal part of calyptra, is known as vaginula (Fig. 10.10B). The foot remains embedded in the vaginula
of the sporophyte.
(iii) Dehiscence of Capsule The capsule dehisces by a special explosive mechanism, usually on a
bright sunny day. This mechanism is also known as air-gun mechanism. Due to heat of the sunny day,
the columella and wall of the capsule become dry and get shrivelled. This promotes the development
of an air space under the spore sac, and the capsule also changes its shape from spherical to cylindrical.
The air present inside the capsule gets compressed, and a pressure also develops inside the capsule.
Due to this, the operculum breaks off at the annulus. The spore sac ruptures and the spores are blown
to a height of several centimetres.
160 � Bryophyta
Fig. 10.10 Sphagnum. A, A female branch bearing perichae al leaves and sporophyte; B, LS of sporophyte
(e) Young Gametophyte The haploid spore develops into a young gametophyte, and is thus the first
cell of the gametophytic generation. The spores, when young, are arranged tetrahedrally. Each spore
varies from 15 to 40 mm in diameter. They are uninucleate bodies, and their wall is made up of a thin
inner endospore and a rough or papillose outer exospore. The spores of Sphagnum remain viable for
as much as 4–6 months. In suitable conditions, the spores may germinate even within 2–3 days after
dispersal.
At the time of germination, the exospore ruptures along the triradiate ridge, and the endospore comes
out in the form of a germ tube (Fig. 10.11A-B). The germ tube divides by transverse divisions to form
a green filament of 2-4 cells. An apical cell with two cutting faces develops in the uppermost cell of
this filament due to two oblique vertical divisions (Fig. 10.11C-D). A plate-like multicellular, thallus-
like structure develops due to the activity of the apical cell. This is known as primary protonema
(Fig. 10.11E). Some marginal cells of this green primary protonema become meristematic and form
secondary protonema, which contain rhizoids and leafy buds (Fig. 10.11F). It is these leafy buds
which develop into mature gametophore of Sphagnum. Secondary protonema does not develop in
many species of Sphagnum, and in such conditions, the leafy gametophores develop from the primary
protonema only.
Fig. 10.11 A-F Sphagnum. Successive stages of spore germina on and forma on of young gametophy c plant
AFFINITIES OF SPHAGNUM 10.3

Sphagnum is unique among bryophytes because on one hand it shows resemblances with Hepaticopsida,
Anthocerotopsida and also with Bryopsida but, on the other hand, it has some unique features of its
own. Some, therefore, consider Sphagnum as a synthetic group. A glimpse of all such resemblances
and unique characters is presented here.
10.3.1 Resemblances of Sphagnum with Hepa copsida

1. Thalloid protonema of Sphagnum (Fig. 10.11E) resembles with that of several liverworts
including acrogynous Jungermanniales.
2. Growth in thalloid protonema of Sphagnum occurs by an apical cell with two cutting faces, as
in some acrogynous Jungermanniales.
3. Mechanism of dehiscence of antheridium of Sphagnum resembles with that of Porella and
some other acrogynous Jungermanniales.
4. The mode of development, position and structure of archegonium of Sphagnum is also same
as that of many acrogynous Jungermanniales.
162 � Bryophyta
10.3.2 Resemblances of Sphagnum with Anthocerotopsida

1. Apical growth is not shown by young sporogonium of Sphagnum and the same is true also in
hornworts.
2. The entire endothecium in Sphagnum develops into sterile columella, as is also the case in
Anthocerotopsida.
3. Archesporium in Sphagnum is amphithecial in origin and the same is the case in
Anthocerotopsida.
4. Wall of the capsule of both Sphagnum and Anthocerotopsida contains several chlorophyll-
containing cells and is thus photosynthetic in nature.
5. Sporophyte in both Sphagnum and hornworts is differentiated into a large bulbous foot, an
ill-developed seta and a well-developed capsule.
10.3.3 Resemblances of Sphagnum with Mosses

1. Presence of erect, leafy gametophores in both.
2. Presence of multicellular rhizoids in both.
3. Presence of oblique septa in the rhizoids of both.
4. Resemblance of leaves of Sphagnum and Leucobryum in possessing alternate living green
cells and dead hyaline cells.
5. Development of antheridium of Sphagnum resembles with that of several mosses.
6. Presence of stalked archegonium in both.
7. Presence of a massive venter in the archegonia of both.
8. Dehiscence of capsule with the help of a lid-like operculum in both Sphagnum and mosses.
10.3.4 Unique Characters of Sphagnum

1. Absence of rhizoids in the adult gametophore.
2. Development of branches in tufts, and that too from the axil of every fourth leaf.
3. Absence of midrib in the leaves.
4. Cortex of mature stem is made up of dead, empty and colourless cells.
5. Several spiral thickenings develop in the dead cells of the cortex.
6. Cortex cells have large oval pores on their walls in many species of Sphagnum.
7. Dead empty cells of cortex absorb water and thus behave like velamen of orchid roots.
8. Flask-shaped retort cells develop in the cortex of side branches of Sphagnum.
9. In spite of being a hydrophytic plant, Sphagnum shows several typical xerophytic characters.
10. The cell walls of this unique bryophytic plant contain some organic substance of colloidal
nature, due to which it absorbs bases and releases acids. Water is, therefore, highly acidic
where Sphagnum grows.
ANDREAEALES 10.4
1. Andreaeales includes the mosses containing a capsule, which resembles a Chinese lantern,
hence called lantern mosses.
2. Since these mosses grow exclusively on siliceous rocks, these are also called granite
mosses.
3. Gametophores are small, quite brittle, dark brown or reddish in colour.
4. The conducting strand is absent in the stem.
5. The perichaetial leaves are quite large, stiff, erect and convolute.
6. The endothecium gives rise to archesporium and columella.
7. A dome-shaped spore sac overarches the columella.
8. Capsule wall lacks spongy photosynthetic tissue.
9. Foot is well-developed, and seta is very short or ill-developed or even may be absent.
10. The mature capsule gets elevated on a specialised gametophytic structure, called
pseudopodium.
11. Dehiscence of capsule takes place by four longitudinal slits, which may sometimes be as many
as ten in number.
12. The protonema is a thalloid and parenchymatous body.
Andreaeales has a single family Andreaeaceae, which includes genera like Andreaea, Neuroloma,
Andreaeobryum and Acroschisma. Some details of the life history of Andreaea are given here.
ANDREAEA 10.5
Andreaea is worldwide in distribution and prefers to grow on siliceous rocky substratum, specially
on exposed rocks in high mountain tops of tropics and colder temperate regions. It grows well in
extremely dry situations on noncalcareous granite rocks, and that is why it has been commonly named
“granite moss”. Five species of Andreaea have been found from different regions of eastern and western
Himalayas. These include A. commutata, A. densifolia, A. indica, A. kashyapii and the most common
cosmopolitan species, i.e., A. rupestris.
The gametophores of Andreaea (Fig. 10.12 A) are green only when young, but even slightly mature
plants are deeply pigmented being orange to deep purple or dark brown to even black. Plants are small-
sized, reaching as much as 2 cm or only slightly more. They form irregular branching to provide a look
of dense compact tuft. The leaves are spirally arranged on the stem, and tend to be imbricate when
dry but divergent when moist. They vary in shape from subulate to ovate and their margin is entire or
sometimes toothed. They are usually unistratose, with or without costa in different species.
Plants grow prostrate along the rock surface. Many multicellular and cylindrical rhizoids develop
from the lower parts of the stem. They usually penetrate into the rock crevices. The stem is very brittle
in dry conditions and can divide easily into fragments, from which new plants may develop, thus
proving to be a method of vegetative propagation.
In Andreaea, the stem grows by means of an apical cell with three cutting faces, and therefore three
rows of leaves are present.
Anatomically, the stem is almost uniform in structure and shows no differentiation into cortex and
central conducting strands (Fig. 10.12B).
Majority of the Andreaea species are monoecious. The sex organs (antheridia and archegonia) are
present on separate branches and are terminal in position. Some species (e.g. A. nivalis) are dioecious.
Growth of the branch containing these sex organs stops after initiation of the sex organs because the
apical cell is responsible for sex-organ formation. The last-formed segments of the apical cell form
164 � Bryophyta
other antheridia or archegonia. Perigonial leaves resemble the vegetative leaves while perichaetial
leaves are bigger than that of remaining leaves. It has also been observed that sometimes a new apical
cell differentiates near the base of the perichaetial leaves. This apical cell develops into new branches
known as innovation organs.
Development of both the sex organs (antheridia and archegonia) follows the same pattern as in
Funaria, and is described in detail under articles. 10.7.7(2) and 10.7.7(6). Antheridia are, however,
ellipsoidal bodies, each with a long stalk which is usually uniseriate and sometimes biseriate (Fig.
10.12C). The archegonium is a short-stalked, flask-shaped body (Fig. 10.12 D) with a long and broad
neck and a slightly globular venter. The first archegonium develops directly from the apical cell of the
female branch.
Fig. 10.12 Andreaea rupestris. A, Part of gametophore (enlarged) with sporophyte; B, TS stem; C, LS apex of
male plant showing antheridia; D, An archegonium; E, Young sporophyte showing differen a on
of archesporium and foot; F, TS young sporophyte showing differen a on of jacket layers,
archesporium and columella
The zygote divides first by a transverse division to form a bicelled embryo. Its lower cell divides
irregularly to form a foot, and the upper cell divides and differentiates an apical cell with two cutting
faces. The foot is an unorganised mass of cells (Fig. 10.12E). The apical cell derived from the upper
cell cuts several segments on each side to form a multicellular body which divides periclinally to form
an outer amphithecium and inner endothecium. A three- to eight-celled thick jacket of the capsule in
derived from the amphithecium. The endothecium again divides periclinally to form two layers, or the
inner endothecium matures into columella (Fig. 10.12F).
The capsule is an elliptical body, and the sporophyte lacks seta (Fi. 10.13A). Some gametophytic
cells elongate and push the capsule through the perichaetial leaves and form a long, leafless stalk
known as pseudopodium (Fig. 10.13B). The jacket of the capsule gets heavily impregnated with black
or dark-brown pigments. Capsule lacks peristome and also the operculum.
Mature sporophyte of Andreaea consists of a well-developed foot and a small capsule. It lacks
structures like seta, operculum and peristome. Some have opined that seta is suppressed in Andreaea.
The capsule attains a length of only 0.5 mm. The capsule tapers a little towards the apex and base.
It remains covered by a wall of 2-6 layers. The centre of the capsule remains occupied by a club-
shaped columella. The spore sac is dome-shaped and bears only spores. The young sporophyte remains
enclosed by a thick calyptra and perichaetial leaves. The elongating function of seta is taken by a body
of gametophytic tissue called pseudopodium.
Fig. 10.13 A, LS of a sporophyte of Andreaea rupestris; B, A sporophyte showing pseudopodium; C, A spore;

D-F, Development of spore and young gametophyte
166 � Bryophyta
Dehiscence of capsule takes place by four lines of dehiscence. The spores are dispersed through the
slits. The dispersed spores remain viable for a few months.
The spore divides and redivides and earlier few divisions occur within the spore coat (Fig. 10.12C-D).
A multicellular body comes out from the spore coat (Fig. 10.12E), and a well-developed protonema
develops. From this develop the gametophores of Andreaea.
BRYALES 10.6
As mentioned earlier under Article 9.3, Bryales are commonly known as true mosses. This is the
largest order of mosses including more than 675 genera and over 14000 species. Parihar (1987) has
treated Bryales as equivalent to the subclass Bryidae of the class Bryopsida.
1. Plant body is foliose, and leaves of the gametophore usually possess a distinct midrib.
2. The midrib region is more than one cell in thickness.
3. The spore develops into a protonema, which is usually filamentous.
4. The rhizoids are multicellular with oblique septa.
5. The zygote divides transversely into an epibasal and a hypobasal cell, and both usually grow
by a two-sided apical cell.
6. The endothecium develops into columella and archesporium.
7. The columella is well-developed. It usually penetrates the sporogenous layer and reaches up
to the apex of the capsule.
8. In between the wall of the capsule and spore sac is usually present an intercellular space
traversed by many filaments of cells.
9. Differing from Andreaeales, the sporogonium in these mosses is never elevated on the
pseudopodium.
10. A well-developed and elongated seta is present in these mosses.
11. The capsule, when mature, is a very complicate structure, and remains differentiated into
several types of tissues.
12. The capsule opens usually by an operculum.
13. A peristome usually covers the spore cavity.
14. Peristome is hygroscopic in nature and helps in spore dispersal.
FUNARIA 10.7
10.7.1 Systema c Posi on (According to Holmes, 1986)
Division —Bryophyta
Class—Bryopsida
Subclass—Bryidae
Cohort—Eubryiidae
Order—Funariales
Family—Funariaceae
Genus—Funaria

Funaria is one of the most common mosses, distributed widely throughout the world. It includes over
120 species, of which more than 15 species have been reported from India. Funaria hygrometrica is
cosmopolitan and best known of all species of different mosses. It grows on moist grounds in close tufts
and also on damp and shady moist rocks, wells, crevices, tree trunks and other similar surroundings.
10.7.3 Mature Gametophyte

Plant body is gametophytic and foliose, consisting of an erect leafy axis attached to the substratum by
means of rhizoids.
The axis or stem is erect, slender, usually branched and attains a height of 1–4 cm. It remains
covered with many leaves, arranged spirally in 3/8 phyllotaxy, i.e. three complete spirals contain eight
leaves. The leaves are simple, ovate, sessile, green and each possesses a broad membranous base and
a pointed apex (Fig. 10.14 B). A distinct midrib is also present in each mature leaf. The midrib is,
however, not present in very young leaves.
Fig. 10.14 Funaria hygrometrica. A, Plant body with sporophyte; B, A leaf; C, Rhizoids showing oblique septa
168 � Bryophyta
Many rhizoids are present at the base of the gametophores. They are multicellular, slender and
well-branched. Rhizoids contains oblique septa (Fig. 10.14C). They are colourless when young but
mature rhizoids become brown or black-cloloured. The chloroplasts may also develop in the cells of
rhizoids, when they are exposed to sunlight. The main functions of rhizoids are anchoring as well as
absorption.

The growth of the stem takes place by a pyramidal apical cell with three cutting faces. Its three lateral
faces cut off three rows of cells by successive divisions. Each of the so formed cells divides periclinally
into inner daughter segment and an outer segment. The inner daughter segment gives rise to the inner
tissues of the stem while the outer segment contributes to the leaf and the outer tissues of the stem. The
leaves in the young apical part of the axis are thus arranged in three vertical rows.
10.7.5 Anatomy
1. Stem or Axis Internally the mature axis is divided into epidermis, cortex and central strand
(Fig. 10.15A, B).
Epidermis is the outermost layer made up of tangentially elongated cells filled with chloroplasts.
Pores or stomata are absent.
Cortex is the well-developed, parenchymatous region of the axis present just inner to the epidermis
and extends up to the central strand or central cylinder. In the young axis, the cells of the cortex also
contain chloroplasts, which disappear in the mature stems. In mature stems the cells of the outer layers
of cortex may also become thick-walled and somewhat reddish-brown, while the cells of the inner
cortex are thin-walled. Isolated small leaf traces may also be seen near the periphery.
Central strand or central cylinder is made up of long, narrow, colourless, thin-walled cells. They
lack protoplasm and are thus dead cells. They help in conduction of water and other nutrients.
Fig. 10.15 Funaria hygrometrica. A, T.S. of a gametophore at a level of junc on of a leaf with the stem;
B, TS of stem and leaf cut at a slightly higher level
2. Leaf
Internally, the leaf is very simple in structure. Its lamina or wings are composed of a single layer of
large thin-walled cells. They remain filled with chloroplasts, and thus represent the main photosyn-
thetic tissue of the gametophore. In the centre of the lamina is present a midrib. It is made up of small
strands of narrow and thick cells. The cells of the central strand help in conduction. Hairs or stomata
are absent on the leaf.

Funaria reproduces vegetatively by various methods including gemmae, bulbils, primary protonema,
secondary protonema and apospory.
1. Gemmae
When conditions are unfavourable, some of the terminal cells of protonema form green multicellular bod-
ies called gemmae. They are both transversely as well as vertically septate. On being detached from the
plant, and if the conditions are favourable, each gemma germinates into a new gametophore of Funaria.
2. Bulbils
Gemmae-like structures developing on rhizoids are known as bulbils. They lack chloroplasts and are
thus nongreen structures. On being detached, each bulbil germinates into a new plant.
3. Primary Protonema
Spore germinates into a branched filamentous structure known as primary protonema. Accidentally
or due to death and decay of some of its cells, the primary protonema gets broken into small fragments.
Buds develop on such fragmented parts of the primary protonema. Each of these buds develops into
gametophore of Funaria.
4. Secondary Protonema
The protonema formed on the injured stems, leaves, reproductive parts or even rhizoids, are known as
secondary protonema. Buds also develop on secondary protonema, and each such bud matures into a
new foliose gametophore.
5. Apospory
Sometimes, diploid gametophores are produced from the vegetative cells of the sporophyte of Funaria,
that is, without the production of spores. This phenomenon is known as apospory. Such gametophores
resemble normal haploid gametophores in appearance but they have diploid cells. Such plants also
produce gametes but they are also diploid. On being fused with the gametes of the opposite sex, the
resultants are 4n zygotes and produce sterile sporophytes.

Funaria is a monoecious and autoecious moss, i.e. male and female reproductive structures develop on
different branches of the same plant. Usually, the female or archegonial branches are longer than that
of male or antheridial branches.
170 � Bryophyta
1. Antheridial Branches
The main axis becomes expanded at the apex, and bears groups of antheridia (Fig. 10.16A) in different
stages of their development. A rosette of spreading leaves also surround the groups of antheridia, and these
are known as perichaetial leaves. Many multicellular, long, capitate and green hairs, intermingled with an-
theridia and perichaetial leaves are also present. These are known as paraphyses. Being green, they contain
chloroplasts, and help in photosynthesis. Antheridia are also protected by these paraphyses.
Fig. 10.16 Funaria. A, LS of an antheridial branch; B, A single antheridium
2. Development of Antheridium
The development of antheridium starts from a superficial antheridial initial (Fig. 10.17A) situated at
the apex of the antheridial branch. This initial soon enlarges, becomes papillate and divides to form
an outer cell and a basal cell (Fig. 10.17B). The basal cell forms the lower part of the stalk of the
antheridium which remains embedded in the tissue. The outer cell divides and redivides transversely
to form a small 2-4 celled filament (Fig. 10.17C). The uppermost cell of this filament divides by two
vertical intersecting divisions, resulting into an apical cell with two cutting faces (Fig. 10.17D). This
apical cell cuts off segments in a regular sequence in two rows (Fig. 10.17E). Few upper cells of this
filament divide diagonally by vertical divisions, each forming two unequal cells, of which the smaller
peripheral cell functions as the first jacket cell (Fig. 10.17F) and the larger daughter cell divides in a
similar fashion to form second jacket initial on the outer side and primary androgonial cells on the in-
ner side (Fig. 10.17G). Two jacket initials and one primary androgonial cell are thus formed by each
cell of the filament. Many anticlinal divisions in the jacket initial give rise to a single-layered jacket of
the antheridium (Fig. 10.17H). Jacket cells, when young, are green in colour because of the presence
of chloroplasts. An operculum is formed by the apical cell of the filament (Fig. 10.17I). Repeated
divisions in the primary androgonial cell make it a multicellular mass of cells which develop into an-
drogonial cells (Fig. 10.17I). Further division in each androgonial cell results in the formation of two
androcytes. Each of the androcytes differentiates into a single, uninucleate and biflagellate anthero-
zoid (Fig. 10.17 K).
3. Mature Antheridium
The mature antheridium (Fig. 10.17 J, K) possesses a long multicellular stalk and a dark-coloured
club-shaped body. The mass of androcytes remains enclosed in the body by a single-layered antheridial
jacket (Fig. 10.17J). A comparatively larger hyaline cell of the operculum is present at the tip of the
jacket layer. Sometimes the operculum consists of two cells (Fig. 10.17 J).
Fig. 10.17 A-K. Funaria hygrometrica. Development of antheridium

172 � Bryophyta
4. Dehiscence of Antheridium
Fully mature antheridium dehisces when it comes in contact with water. The operculum cell absorbs
water, becomes mucilaginous and swells up. Due to pressure, it bursts forming a pore-like structure at
the distal end of the antheridium. The mass of androcytes surrounded by the viscous fluid oozes out
through this pore. Contraction of the antheridial wall also helps in releasing the mass of androcytes.
The androcytes spread out in the form of a thin film, their membranous walls dissolve in water, this all
finally liberates the spirally coiled biflagellate antherozoids.
5. Archegonial Branches
The female or archegonial branches in Funaria usually develop laterally at the base of the male branch.
At the apex of these branches develop archegonia (Fig. 10.18) surrounded by many leaves known as
perichaetial leaves. Along with archegonia and perichaetial leaves are also present several paraphy-
ses on the archegonial branch. The cells of both perichaetial leaves and paraphyses remain filled with
several chloroplasts.
Fig. 10.18 Funaria hygrometrica. Longitudinal sec on of an archegonial branch bearing archegonia
6. Development of Archegonium
It starts from a superficial cell, known as archegonial initial (Fig. 10.19A), which gets differentiated
at the tip of the archegonial branch. It first divides transversely to form a lower cell and an upper cell
(Fig. 10.19B). Repeated divisions in the lower cell results in the formation of the basal part of the
archegonium which remains embedded in the tissues of the archegonial branch. Upper cell starts func-
tioning as archegonial mother cell and divides by two intersecting oblique walls to form an apical
cell with two cutting faces (Fig. 10.19C). This apical cell cuts many (4–8) segments alternately on both
sides and forms the archegonial stalk (Fig. 10.19D). The apical cell now divides by three intersecting
oblique divisions to form three peripheral cells surrounding an axial cell (Fig. 10.19E). Each periph-
eral cell divides anticlinally and forms a single-layered jacket of the venter. By further divisions, it
becomes bilayered. The derivatives of the axial cell develop into the neck of the archegonium.
Fig. 10.19 A-I. Funaria hygrometrica showing successive stages of the development of archegonium
The axial cell (Fig. 10.19E) divides transversely into a primary cover cell and an inner central
cell (Fig. 10.19 F). The primary cover cell now behaves as an apical cell with four cutting faces. The
174 � Bryophyta
derivatives of three of these faces form the jacket of the neck whereas the fourth basal face forms a row
of neck canal cells. The central cell divides transversely into an outer primary neck canal cell and
inner primary venter cell (Fig. 10.19G). The primary neck canal cell divides transversely to form a
row of neck canal cells and all these occupy the median and lower part of the neck. In Funaria, the neck
canal cells, therefore, have double origin, i.e. those present in the median and lower part of the neck
originate from the primary neck canal cells while those present in the upper part of the neck originate
from the basal derivatives of the cover cell. The primary venter cell divides transversely to form a
venter canal cell and an egg (Fig. 10.19 H,I).
7. Mature Archegonium
Mature archegonium (Fig. 10.19I) consists of a well-developed stalk, a swollen venter and an elon-
gated neck. The venter region is surrounded by a bilayered archegonial jacket while in the neck region,
it is single-layered. The neck contains a row of 6-9 or more neck canal cells, and the venter contains a
venter canal cell and an egg cell (Fig. 10.19 H, I).
8. Fer liza on
Fertilization is facilitated by rain or dew water. Biflagellate antherozoids are set free during the process
of dehiscence of antheridium. On the other hand, the neck canal cells and the venter canal cell of the
archegonium disintegrate and degenerate, and form a mucilaginous mass. This mass absorbs water,
and due to this the cover cells present at the top of the neck of the archegonium are forced apart. A free
passage for the entry of antheridium is thus formed inside the archegonium. Dew or rainwater helps
in the transfer of antherzoids from antheridial head to the archegonial head. Water drops containing
antherozoids easily tickle down from antheridial to archegonial head because the archegonial heads are
usually at the lower level than the antheridial heads. Water current helps in transfer of antherozoids
from antheridial heads to archegonial heads in submerged species of Funaria.
The antherozoids enter into the archegonium due to the chemotactic imfluence of the mucilaginous
substances of the neck. Many antherozoids enter inside the archegonial neck, but ultimately only one
of them fuses with the egg and forms a diploid zygote.
10.7.8 Sporophyte
Fusion of antherozoids and an egg in the process of fertilization results in the formation of a diploid
zygote, which is the first cell of the sporophytic generation. A wall is secreted around the zygote. It also
increases in size and fills the cavity of the venter (Fig. 10.20A).
1. Development of Sporophyte
The diploid zygote first divides by a transverse division into an upper epibasal cell and a lower
hypobasal cell (Fig. 10.20B). Both of these cells divide further by two oblique intersecting walls,
resulting in the formation of a two-sided apical cell in both epibasal cell as well as hypobasal cell
(Fig. 10.20C). Two growing points, represented by two apical cells, are thus present in the young
embryo at this stage. Each of these apical cells has two cutting faces. The epibasal cell (Fig. 10.20B)
and its derivatives give rise to capsule and a part of the seta while the hypobasal cell and its derivatives
give rise to foot and remaining part of the seta. The cells of the wall of the venter divide and redivide to
form a calyptra, which is a protective covering and covers the capsule till it matures.
Fig. 10.20A-E Funaria. A, Mature zygote inside the archegonium; B-E, Early stages of the
development of sporophyte
(a) Development of Capsule

Study of the serial transverse sections of the young sporophyte can provide us a clear picture of the
development of the capsule. If a transverse section just below the apical cell is cut, its two derivatives
form an almost spherical segment (Fig. 10.21 A). These two derivatives are one each from two faces
of the epibasal apical cell. A vertical division in both these derivatives forms the quadrant stage of the
embryo. (Fig. 10.21B). In this quadrant embryo, each cell divides by an anticlinal division in such a
way that two such cells are resulted, of which one is smaller and almost triangular while the other one
is larger and more or less rectangular in shape (Fig. 10.21C). This stage contains eight cells of the
embryo, in which four are triangular cells while the remaining four are rectangular cells, and these are
alternately arranged. Each rectangular cell divides by a periclinal division. In this way, at this stage
eight peripheral cells, which form amphithecium, surround the centrally-located four cells of the en-
dothecium (Fig. 10.21D). These two (amphithecium and endothecium) are fundamental embryonic
layers of the sporophyte. Cells of these two layers divide in a definite pattern and form three regions of
the capsule, i.e. apophysis, theca and operculum, from base towards top.
(i) Early Fate of Amphithecium
The amphithecial cells (Fig. 10.21D) divide periclinally and develop into two concentric rings made up
of 8 cells each. The inner ring is designated as the first ring (Fig. 10.21 E). In the outer ring, the cells
first divide anticlinally and then periclinally forming two concentric rings of 16 cells each. The inner
one of these two concentric rings is designated as the second ring (Fig. 10.21G). The cells of the outer
ring divide first anticlinally resulting into 32 cells and then periclinally forming two concentric rings of
32 cells each. The inner ring of 32 cells is the third ring, and the outer ring divides again periclinally,
thus resulting into two rings of 32 cells each (Fig. 10.21G). Of these two rings of 32 cells each, the
inner one is designated as fourth ring and the outer one is known as the fifth ring (Fig. 10.21H, I). An
additional sixth ring is formed in the opercular region.
176 � Bryophyta
(ii) Early Fate of Endothecium As mentioned above, the endothecium is present in the form of a central
core of four cells (Fig. 10.21 D,E). These cells repeat the same pattern of divisions that differentiates
the amphithecium and endothecium. Due to this, a group of four endothecial cells is differentiated by
eight peripheral cells (Fig. 10.21F,G). Two subsequent divisions in the endothecial cells give rise to 16
cells, which form columella. Eight peripheral cells divide first by radial wall and then by a periclinal
division. Two layers of 16 cells each are thus differentiated. The inner layer, which is adjacent to the
columella, matures into the inner spore sac, while the cells of the outer layer divide again by radial
walls to form the single-layered archesporium. The single-layered archesporium divides periclinally
and soon becomes double-layered. All archesporial cells are fertile and behave as spore mother cells.
Reduction division in each archesporial cell results in the formation of four haploid spores.
(iii) Fate of the Rings in the Development of Operculum, Theca and Apophyses in the Young Capsule
• Fate of Endothecium in the Operculum It develops into central parenchymatous region.
The first ring contributes to outer peristomial layer. The second ring gives rise to the middle
peristomial layer. The third ring contributes to the inner peristomial layer.
Fig. 10.21 A-I Funaria. Stages of the development of sporophyte, including differen a on of
amphithecium and endothecium (A-D) and forma on of five rings of
amphithecial cells in the region of capsule (E-I)
• Fate of Amphithecium in the Operculum The fourth and fifth rings contribute to the inner
layer of the operculum. The sixth ring, present only in the operculum region, contributes to the
epidermis of the operculum and annulus.
• Fate of Endothecium in the Theca Four central cells contribute to the columella. Inner layer
of 16 peripheral archesporial cells form the inner wall layer of the spore sac. Peripheral cells
form the outer layer. The outer layer of 16 peripheral archesporial cells form the archesporium
which forms spore mother cells.
• Fate of Amphithecium in the Theca The first ring contributes to the outer wall of the spore
sac. The second ring gives rise to trabeculae. The third ring forms chlorenchymatous layers.
The fourth ring forms the hypodermis. The fifth ring contributes to the epidermis.
• Fate of Endothecium in the Apophysis It contributes to the central conducting strand. Four
inner layers form spongy tissue around the conducting strand.
• Fate of Amphithecium in the Apophysis The outermost fifth layer contributes to the epidermis
of the apophysis region of the young capsule.
3. Structure of Mature Sporophyte

The sporophyte (Fig. 10.22A) is differentiated into foot, seta and capsule. The foot is a small, conical
body embedded in the tip of the archegonial branch. The seta is very long, more or less twisted and a
stalk-like structure which supports the capsule at its tip.
Internally, the seta is differentiated into an outermost layer of epidermis, enclosing somewhat
thick-walled cells of the cortex and a centrally-located central conducting strand (Fig. 10.23) made
up of thin-walled elongated cells. Functions of seta include (i) transport of water and nutrients to the
capsule, and (ii) to provide mechanical support to the capsule.
The capsule is erect and green, when young, but at maturity it becomes curved and bright orange
coloured. The capsule can be divided into three major regions (Fig. 10.22B), viz. (i) apophysis, (ii)
theca proper, and (iii) apical region.
(a) Apophysis The apophysis is the basal sterile part of the capsule which connects the capsule with
the seta of the sporophyte. It is made up of the central conducting strand, which extends down into
the seta. Around the central strand are present loosely arranged chlorophyll-containing cells. On its
outermost side is present a layer of epidermis, the continuity of which is broken by stomata. Because
of the presence of chlorophyllous cells, the apophysis performs the function of photosynthesis. The
Funaria sporophyte is, therefore, partially dependent on the gametophyte.
(b) Theca Proper Theca is the middle part of the capsule, present in between the apical region and
apophysis. It is the fertile part of the capsule, and contains various parts, including columella, spore
sac, air space and capsule wall.
• Columella It is the centrally-located, cylindrical, pith-like part of the theca made up of
parenchymatous cells. Its distal end projects into the operculum. At its base, the columella
remains connected with the apophysis.
• Spore Sacs Two elongated spore sacs surround the columella. On the outer side, each spore
sac remains covered by a 3–4 layered wall. The wall present on the inner side of the spore sac
178 � Bryophyta
is single-layered. Each spore sac contains many spore mother cells, when young. At maturity,
each spore mother cell forms four haploid spores. Funaria lacks elaters.
Fig. 10.22 Funaria. A, A gametophyte with sporophyte; B, LS capsule
• Air Space Outside each spore sac is present a large air space or air cavity. It remains traversed
by many filaments which are green, delicate and contain chlorophyll-containing cells. These
filaments are known as trabeculae.
• Capsule Wall The wall of the capsule is made up of many layers of thin-walled parenchymatous
cells. The epidermis, followed by 2-3 layered hypodermis, is present on the outermost side
of the capsule wall. Inner to the parenchymatous hypodermis are present 2–3 layers of
chlorophyllous cells, which enclose many intercellular spaces. Through trabeculae, the
innermost layer of the capsule wall remains connected with the outer wall of the spore sac.
Fig. 10.23 Funaria. TS of seta
(c) Apical Region

Two main parts of the apical region of the capsule are operculum and peristome (Fig. 10.22B). A
constriction is present at the juncture of the theca and apical regions. A rim or diaphragm is present
immediately below this constriction. The rim, made up of 2–3 layers of radially elongated cells,
demarcates the upper limit of the theca.
• Operculum The operculum is a lid-like conical structure (Fig. 10.22B) which closes the mouth
of the capsule. It is made up of 2–3 layers of thin-walled cells. The cells of its outermost layer
have thickened outer walls. At the broader lower end of the opercular lid, a few layers form a
ring of large, well-developed cells, which represent the annulus. The annulus is composed of
thin-walled cells. It helps in the dehiscence of the capsule.
• Peristome Immediately below the operculum lies the peristome (Fig. 10.22B). It remains
attached below the edge of the diaphragm. The peristome consists of two rings of long, curved,
triangular teeth-like structures known as peristomial teeth. 16 teeth are present in each ring of
peristome. The outer peristomial teeth are large, thicker and red-coloured structures, while the
inner peristomial teeth are smaller, delicate and colourless structures (Fig. 10.24 A-C). Well-
marked transverse thickening bands are present in the outer peristomial teeth.
4. Dehiscence of Capsule and Dispersal of Spores

Water supply is almost cut off when the sporophyte is fully mature. This results into drying up of all tis-
sues of the capsule. Soon the thin-walled cells of the annulus break and the operculum is thrown away.
In spite of all these conditions, the spores are not dispersed because the mouth of the theca is covered
by the peristome. Being hygroscopic in nature, the outer peristomial teeth bend outwards in dry condi-
tions while the inner peristomial teeth remains as such in their normal position. Due to jerky movement
180 � Bryophyta
of the outer peristomial teeth, the slits present in between inner peristomial teeth become broader, and
through these slits the spores are dispersed. During humid or moist conditions, the outer peristomial
teeth absorb moisture and they show a bending towards the inner side. Due to this, the slits close and
this prevents the escape of spores. Dispersal of spores in Funaria is also facilitated due to twisting and
bending nature of seta of the mature sporophyte.
Fig. 10.24 Funaria. A, A capsule showing peristome; B, Outer peristome in surface view;
C, An outer and an inner peristomial teeth
10.7.9 Young Gametophyte

The dispersed spore germinates (Fig. 10.25A-E) readily to form one or two germ tubes, which elongate to
form a branched filamentous protonema. Initially, the protonema is made up of highly chlorophyllous,
green short cells with transverse septa. This stage of the protonema is known as chloronema. The
chloronema matures into caulonema, which is a shoot-producing protonema. The caulonema is made
up of erect filaments and also prostrate filaments, which comprise green elongated cells with oblique
septa. An initial develops from the cell of prostrate filament, and this initial either develops into a branch
or a gametophore. The branch initial divides by transverse divisions to form a branch but a bud initial
divides by oblique transverse walls to form an apical cell with three-cutting faces. It produces stem and
leaves. From the base of the bud develop the rhizoids, and thus develops a young gametophore.
Fig. 10.25 A-E Funaria, showing germina on of spore (A, B) and successive stages (C-E) of the
forma on of primary protonema, secondary protonema and young gametophore
BUXBAUMIA: A BOTANICAL NOTE 10.8

Systema c Posi on
Class—Bryopsida
Subclass—Bryidae
Cohort—Buxbaumiidae
Order—Buxbaumiales*
Genus—Buxbaumia
Buxbaumia is a microscopic moss, occurring on nutrient-poor soil, decaying logs and other
similar organic substances. It bears very small gametophytes and a bug-like dorsiventrally flattened
sporophyte. The sporophytes attain a length of about 2 cm, and only after their formation the
microscopic gametophytes can be noticed easily. It is a moss, mainly of temperate regions of the
northern hemisphere, and about 15 of its species have so far been reported. Udar et al. (1971) reported
Buxbaumia himalayensis from India.
The perigonium of this microscopic moss is the most reduced gametophyte known in bryophytes
(The leaves or bracts surrounding the sex organs of bryophytes, specially those around the antheridia,
are termed perigonia). The male gametophyte (Fig. 10.26 A,B) comprises a single unistratose flap of
tissue which surrounds a single, spherical, short-stalked antheridium. In the female gametophyte, the
perichaetium is also very small, made up of 3–4 unistratose perichaetial leaves (Fig. 10.26C), which
enclose one or two archegonia and a few, very small paraphyses.
*Buxbaumiales are commonly called bug mosses.

182 � Bryophyta
Fig. 10.26 A-E, Buxbaumia aphyllla. A, A male gametophyte; B, Ver cal sec on of a male
gametophore showing single antheridium; C, A female gametophyte of B. aphylla;
D, Sporophyte; E, LS of capsule drawn diagramma cally
The sporophyte (Fig. 10.26D) comprises foot, seta and capsule. The foot is a bulbous body
embedded within the upper part of the perichaetium. Ths sporophyte, when young, is a green body
containing a small conical calyptra at the top of the seta. The seta is a papillose body containing red
pigments in its outer thick-walled cells. A small conducting strand is present in the centre of the seta.
It is as long as 1.5 cm. The capsule is oval in shape and contains a short conical operculum and a
broad expanded apophysis (Fig. 10.26D). A characteristic feature of the capsule of Buxbaumia is that
it is oriented obliquely at the apex of the seta in a way that one face appears a flattened structure. The
capsule remains surrounded by a multistratose jacket. The stomata are present only in the region of
apophysis. A well-developed cylinder of intercellular space is present in between the jacket and the
sporogenous tissue. The mature capsule is a glossy, chestnut-coloured structure attaining a length of
as much as 0.5 cm.
In Buxbaumia, the peristome consists of a most complex structure found among mosses. Each
peristomial ring contains 32 teeth, and there may be as many as five concentric rings in the peristome.
The teeth are shortest in the outermost ring and gradually become longer in each outer row. The outer
teeth are termed exostome. Each tooth of the peristome is an individual and isolated body. In all
species, there is present a central truncated core of fused peristomial teeth. It is termed as endostome.
The peristomial teeth in Buxbaumia are not hygroscopic, and due to this it is not confirmed whether or
not they help in spore dispersal.
Germination of spore starts by producing a small germ tube made up of green cells representing
chloronema. From the chloronema develops caulonema. A bud is produced on the caulonema, which
can develop either into a female gametophyte or male gametophyte.
TETRAPHIS: A BOTANICAL NOTE 10.9

Systema c Posi on
Class—Bryopsida
Subclass—Bryidae
Cohort—Eubryiidae
Order—Tetraphidales
Family—Tetraphidaceae
Genus—Tetraphis
Tetraphis is one of the two genera of the family Tetraphidaceae. The another genus of the family
is Tetradontium. Tetraphis is important because of some unique features of its sporophyte. Due to the
presence of four teeth in its peristome, Tetraphis is known as four-toothed moss.
Tetraphis occurs commonly in coniferous forests distributed widely in the northern hemisphere.
Plants are small-sized (less than 1 cm in length) and grow on peat banks, decomposing wood and other
similar surroundings.
The gametophytic plant body remains attached to the substratum by means of uniseriate rhizoids.
Leaves are very small on the lower side of the axis but large on its upper portion. They are ovate, costate
and bright green in colour (Fig. 10.27A). At the top of the young gametophyte is present a cup-like
cluster of short blunt leaves enclosing several gemmae (Fig. 10.27B). Each gemma is a multicellular,
lens-shaped body with a long stalk (Fig. 10.27C). It serves as a major method of vegetative propagation
in Tetraphis.
Anatomically, a single-layered epidermis surrounds a well-developed cortex, differentiated usually
into an outer cortex of thick-walled cells and an inner cortex of broad thin-walled cells. A narrow strand
of centrally-located, small-sized cells serves the purpose of conducting strands, which are known as
hydroids.
Tetraphis is a monoecious moss. The sex organs (antheridia and archegonia) are present terminally
on separate branches in groups of 8–10 or more, and remain surrounded by a cluster of elongate
perichaetial or perigonial leaves, along with many filamentous paraphyses. After the formation of sex
organs the growth of the plant stops.
The sporophyte is green and chlorophyllous when young. It consists of foot, seta and capsule. The
foot is tapered, penetrates the apex of the gametophyte and serves the functions of anchorage and
absorption. The seta is very long and contains a central conducting strand. It is quite rigid, and its
rigidity is due to the presence of thick-walled stereids present outside the conducting strand. The jacket
of the capsule in Tetraphis is derived from the amphithecium while the other remaining tissues of the
capsule are derived from the endothecium. The columella is centrally-located in the capsule in the form
184 � Bryophyta
of a multicellular central cylinder. A sporogenous layer surrounds the columella. The apical portion
of the jacket of the sporophyte develops into the operculum. The peristomial teeth are four in number
(Fig. 10.27E), and each one is multicellular and wedge-shaped. The calyptra persists till the capsule is
fully mature.
The jacket of the mature capsule contains elongate and thick-walled cells in its outermost layer, in
which stomata are absent. At the time of dehiscence, the capsule dries. Due to dryness, the operculum
is shed, the peristomial teeth are exposed and the spores are then dispersed. Twisting and untwisting
of seta also help in spore dispersal. Resembling Funaria, the peristomial teeth in Tetraphis are also
hygroscopic in nature. In moist conditions, they move inwards but they get apart in dry conditions.
Dispersal of spores is checked in excessive moist weather.
Fig. 10.27 A-E Tetraphis pellucida. A, An enlarged gametophyte of T. pellucida with a sporophyte;
B, Upper part of a plant bearing gemma cup and gemmae; C, An enlarged gemma;
D, T.S. stem of T. pellucida; E, A capsule with peristomial teeth
The spores are unicellular. They germinate by producing a well-branched and green filamentous
protonema, called chloronema. It gives rise to perpendicular, green protonemal flaps, which are
chlorophyllous, unistratose and strap-shaped bodies. Near the base of the protonemal flaps originate the
shoot buds, from which leafy gametophytes are formed. Tetraphis resembles Sphagnum in possessing
a flat, plate-like protonema.
ARCHIDIUM: A BOTANICAL NOTE 10.10

Archidium is the monotypic genus of the only family Archidiaceae of order Archidiales of Bryopsida.
It is represented by more than 25 species, and due to the unusually large-sized spores, these mosses are
commonly known as large-spored mosses.
Archidium is distributed widely in temperate regions of the globe. The gametophores (Fig. 10.28A)
are very small-sized, attaining a height of about 2 mm to 2 cm. Plants usually occur on clay, sand or
gravel and moist soils of somewhat disturbed habitats. The axis or stem is erect and remains attached
to the substratum by many rhizoids. The leaves remain spirally arranged on the stem. The leaves are
usually clasping and of varying shapes, but generally they are ovate to slightly lanceolate. All leaves
are costate and the cells of multistratose costa are uniformly thick-walled. The plants are annual or
perennial. Formation of innovations is quite common.
Fig. 10.28 A-D, Archidium. A, An enlarged gametophyte; B, A gametophyte bearing antheridia;

C, Sporophyte with a massive foot and no seta; D, Sporophyte bearing large-sized spores
186 � Bryophyta
Anatomically, the axis comprises of mainly simple, large parenchymatous cells. The cells of the
outer side are quite large and those of inner central regions are thin-walled and somewhat compact.
Archidium gametophytes are monoecious. The antheridia (Fig. 10.28B), produced at the apex of
branches, remain enclosed by many perigonial leaves. Some filiform paraphyses are also present. The
archegonia also develop at the tips of the branches and remain surrounded by few perichaetial leaves
and ill-developed paraphyses.
The sporophyte of Archidium is quite unique among mosses. Columella is absent in the capsule, and
the seta or stalk of the capsule is also absent or ill-developed. The differentiation of amphithecium is
also unique in Archidium.
The zygote divides first by a transverse division into an epibasal and a hypobasal cell. The hypobasal
cell divides several times to form a multicellular, massive foot (Fig. 10.28). The remaining parts of
the sporophyte develop from the epibasal cell. Development of sporophyte is unique because in a
developing embryo, the four cells of a quadrant are unequal in size. The periclinal divisions form
the amphithecium and endothecium. A dome-shaped intercellular space develops between the
amphithecium and endothecium. The sporogenous tissue is endothecial in origin. The number of spores
in a capsule varies in different species. Sometimes, they are only four in a capsule but in many species
their number reaches up to 60 in a capsule, and in A. winteri, up to 176 spores per capsule have been
reported. The jacket surrounding the capsule is unistratose.
Unusually large-sized spores are present in the capsule of Archidium. They attain a diameter of 50
to 120 microns, and can be seen even with an unaided eye. The spore germination is also unique. A
germinating spore forms a germ tube, developing into a protonema. A quadrant of unequal-sized cells
are seen in this protonema. Of these, one starts forming an apical cell, which divides and redivides to
form a new gametophyte.
POGONATUM: A BOTANICAL NOTE 10.11

Systema c Posi on (According to Holmes, 1986)
Class—Bryopsida
Subclass—Bryidae
Cohort—Polytrichiidae
Order—Polytrichales
Family—Polytrichaceae
Genus—Pogonatum
Plant body of Pogonatum is gametophytic, and each gametophore is differentiated into rhizoids,
axis or stem and leaves (Fig. 10.29A-B). The lowermost rhizomatous part of the stem is stout, stiff and
bears rhizoids. The rhizoids are thick-walled, multicellular and bear oblique septa. The axis is erect,
aerial, well-developed, and bears many leaves. Leaves are well-developed and pale-coloured. Each
leaf is sessile with a sheathing broad base. Its upper part gradually tapers towards the apex and bears a
serrated margin. The leaf has a thick midrib. Parallel, longitudinal, plate-like structures, called lamellae
(Fig. 10.29C) are present on the upper surface of the midrib of the leaves.
Fig. 10.29 A-D. Pogonatum. External features. A, Two female plants; B, A male plant;
C, Upper part of leaf showing lamellae; D, A part cellular of TS axis
Anatomically, the axis (Fig. 10.29D) contains epidermis, cortex, leptoid mantle, stereids and
hydroids. The epidermis is single-layered and consists of slightly thick-walled cells. A wide zone
of cortex is present below the epidermis. The cortex is differentiated into deeply coloured and thick-
walled outer cortex, and a very wide region of thin-walled and parenchymatous inner cortex. A few
thick-walled leaf traces are also present in the thin-walled inner cortex. A well-developed zone of
elongated cells having no starch represent the leptoid mantle inside the inner cortex. The cells of the
leptoid mantle resemble sieve tubes and also contain sieve plate-like structures. These leptoid cells
resemble the leptoid mantle of the axis. A hydrom cylinder is present in the centre of the axis. It is made
up of two types of cells:(i) thick-walled, elongated cells with living contents, called stereids, and (ii)
thick-walled elongated cells without living contents, called hydroids. The hydroids help in conduction
of water.
The male plant (Fig. 10.29B) bears many antheridia in a cluster in its apical part. Antheridia remain
surrounded by specialised leaves called perichaetial leaves. All these together constitute a flower-like
antheridial head (Fig. 10.30A). Many long and multicellular, hair-like paraphyses are also present in
the antheridial head along with a cluster of antheridia (Fig. 10.30B). Each antheridium is a shortly-
stalked, club-shaped structure with multicellular antheridial stalk. A single-layered jacket surrounds
each antheridium.
188 � Bryophyta
The female plant (Fig. 10.29 A,B) bears a cluster of archegonia at its tip (Fig. 10.30C). Along
with multicellular, long, hair-like paraphyses and several perichaetial leaves, the group of archegonia
constitutes the female head. Each archegonium is attached at the apex of the axis by a short multicellular
stalk. It contains a neck and venter. The neck is made up of 6 vertical rows of cells, 2 cover cells and
6–12 or more neck canal cells. The venter consists of a venter canal cell and an egg.
Fig. 10.30 A-E, Pogonatum. A, A detached antheridial head; B, LS antheridial head; C, LS of a

female head; D, An archegonium; E, LS of sporophyte (diagramma c)
The sporophyte, when mature, reveals that it is divisible into foot, seta and a calyptra-covered
capsule (Figs. 10.29 A, 10-30E). The foot is bulbous and seta is very long. The capsule has a long
well-developed lower stalk. It is divisible into a lower stalk, middle fertile region and the uppermost
operculum (Fig. 10.30E). It lacks apophysis. The stalk is parenchymatous and remains surrounded by
green, chlorophyll-filled cells. It appears to be merging with the columella of the capsule. The fertile
region of the capsule shows the following structures:
1. A single-layered epidermis surrounds a many-layered, green region of chlorophyllous cells
followed by a region of air spaces.
2. A region of air spaces surrounds the spore-sac cylinder.
3. The spore-sac cylinder contains archesporial tissue and remains surrounded by a bilayered
spore-sac wall.
4. Archesporial tissue contains many spores at maturity.
5. Yet another air-space region is present inner to the inner spore wall.
6. The inner air space remains connected with a centrally-located parenchymatous columella.
Columella of the capsule passes into the region of operculum and swells in the form of the roof
of capsule called tympanum or epiphragm. The operculum is a beak-shaped or conical structure
above the epiphragm. It remains connected to the capsule by a ring-like diaphragm. A ring of 32 short
peristomial teeth is present above the diaphragm. The peristomial teeth are hygroscopic in nature, and
help in dispersal of spores.
POLYTRICHUM 10.12
10.12.1 Systema c Posi on (According to Holmes, 1986)
Class—Bryopsida
Subclass—Bryidae
Cohort—Polytrichiidae
Order —Polytrichales
Family—Polytrichaceae
Genus—Polytrichum

Represented by about 100 species, Polytrichum is a cosmopolitan genus, but occurs mainly in cool
temperate and tropical regions of the world. It grows commonly along the margins of ponds, lakes, etc.
as well as on sandy grounds, dry woods, damp soil, marshy surroundings, etc. Polytrichum alpinum, P.
commune, P. densifolium and P. juniperinum are some of the common Indian species. P. commune is
cosmopolitan in distribution.

The gametophore of Polytrichum is foliose, moss-like, and differentiated into rhizoids, underground
rhizome, an erect aerial stem and leaves (Fig. 10.31A). The plants attain a height of 20-35 cm or
even more. The rhizoids are numerous, long, thick-walled, multicellular, and with oblique septa.
190 � Bryophyta
Innumerable rhizoids coil around one another to form a twisted or tufted strand and provide mechanical
support to the plant. Water also passes upwards through these tufted strands of rhizoids by external
capillarity. The rhizome is the underground rhizoid-bearing part of the stem. The aerial stem is erect,
unbranched or sometimes branched, leaf-bearing part of the plant. The leaves are usually spirally
arranged on the stem, and are green, brown or even colourless. On the upper part of the rhizome and
on the middle transitional region of the stem, the leaves are arranged in three vertical rows, showing
1/3 type of phyllotaxy, while on the erect leafy shoot the leaves are arranged in a complicated spiral,
showing 3/8 type of phyllotaxy. Each leaf (Fig. 10.31B) contains a membranous colourless sheathing
base and narrows gradually towards the tip. It possesses a midrib and a coarsely toothed margin. The
limb of each leaf is lanceolate to linear lanceolate. On the upper or adaxial surface of the midrib
are present close-set rows of one-celled thick longitudinal plates of green tissue called lamellae. The
midrib appears firm and dark green because of these lamellae.
10.12.4 Anatomy of Rhizome

It is circular or triangular in outline with rounded
corners (Fig. 10.32A). The outermost layer is
the epidermis or piliferous layer, some cells of
which give rise to rhizoids. Below the epidermis
are two to four layers of parenchymatous cortex.
Three sclerenchymatous or parenchymatous
hypodermal strands are present in the three
corners of the cortex, dividing the latter into
three parts. These hypodermal strands form the
radial strands in the centre. The continuity
of the endodermis is broken by these radial
strands. The pericycle is 2–3 layered and
discontinuous because of the radial strands.
The central cylinder is a trilobed structure
with three furrows, and consists of compact
mass of thick-walled tissues. Elongated cells
with very thick walls form the central mass of
this cylinder. These cells are living with oblique
end walls and a small amount of starch, and are
called stereids. The stereids are collectively
called stereom.
Along with the stereids are present certain
other elements of the same diameter but of
different nature. They often fuse together in the
form of bands of two or three cells, separated Fig. 10.31 A, A female plant of Polytrichum
from each other by delicate cellulose walls, commune with a terminal sporophyte;
and function as water-conducting tissue. B, An enlarged leaf
These are individually called hydroids and
collectively called hydrom. In the furrows of the interrupted pericycle are present 6 to 8 polygonal cells
of proteinaceous nature. These cells are individually called leptoids and collectively called leptom. In
between the leptom and hydrom are present some starchy parenchymatous cells called amylom. The terms,
such as stereom, hydrom, leptom and amylom, etc. have been proposed by Tansley and Chick (1901).
Fig. 10.32 Anatomy of Polytrichum commune. A, TS rhizome; B, TS aerial leafy stem; C, TS leaf
192 � Bryophyta
10.12.5 Anatomy of Aerial Leafy Stem

The aerial leafy stem is irregular in outline because of the attachment of leaves (Fig. 10.32 B). The
epidermis is single-layered but not sharply defined. The cortex, present just inner to the epidermis, is
irregular, and consists of compact, elongated and prosenchymatous cells, thus representing the outer
cortex. The inner cortex cells are parenchymatous. Leaf traces are present at different locations of
the cortex. The pericycle is rudimentary and discontinuous. Inner to the pericycle is the region of the
leptom mantle, consisting of cells like that of “sieve tubes”, and thus representing a region like that
of the phloem of higher plants. Amylom layer is situated inner to the leptom mantle. Its cells are
filled with starch. Inner to the amylom layer is the hydrom mantle, the cells of which are thin-walled.
Hydrom cylinder, a region of thick-walled cells, is present in the centre of the stem. It is the water-
conducting tissue of the aerial leafy stem.
10.12.6 Anatomy of Leaf

The leaf is a boat-shaped structure in TS (Fig. 10.32 C). A distinct epidermal layer is present on the
lower or dorsal surface of the leaf. This layer is cuticularised. One or two layers of sclerenchymatous
cells are present just inner to the epidermis. The midrib region is centrally-located and many-celled
thick. The number of cells decreases towards both the sides, forming the wings. A major part of the
leaf is filled by several large parenchymatous cells. Many vertical filaments (lamellae) of chlorophyll-
containing cells are present on the ventral or upper surface of the leaf. Each filament is 5–8 or more
cells in height and contains a papillose or bifurcated terminal cell. Lamellae are the main photosynthetic
regions of the leaf.

Polytrichum plants are usually dioecious, i.e. male and female sex organs develop at the apex of separate
plants.
10.12.8 Antheridial Head

The antheridia are present at the apex of the male gametophore along with several perichaetial leaves
and form a clustrous structure. This clustrous structure looks like an open cup or a flower. Some of
these perichaetial leaves form hair-like structures called paraphyses (Fig. 10.33 A-C). The tips of
some of the paraphyses become a mass of one-celled thick multicellular body. The apical cell of the
shoot is not utilised in the formation of an antheridium. Because of this, the growth of the antheridial
head is not stopped by the development of antheridia. The mature antheridium (Fig. 10.33 C) is a
short-stalked, club-shaped body consisting of several androgonial cells, surrounded by a single-layered
sterile jacket.
10.12.9 Archegonial Head

Terminal clusters of archegonia (Fig. 10.34A-C) develop at the apex of the female gametophore and
form the archegonial head. The archegonia are also surrounded by the perichaetial leaves. Along with
the archegonia are also present many hair-like, multicellular structures. Unlike the antheridial head, the
apical cell of the female gametophore develops into the archegonial initial, and due to this, the growth
of the archegonial head is stopped after the development of archegonia. Each mature archegonium
is a stalked structure (Fig. 25-4C) with a long neck and globular venter. The neck contains several
neck canal cells and remains surrounded by six vertical rows of cells. The venter is multicellular and
contains a venter canal cell and an egg.
Fig. 10.33A-C Details of the antheridial head of Fig. 10.34A-C Details of the archegonial head of
Polytrichum commune. A, A male Polytrichum. A, A female gametophore;
gametophore; B, A part of antheridial B, A part of archegonial head; C, An
head; C, An antheridium along with archegnoium
paraphyses
During the fertilization process, the neck canal cells and venter canal cells disintegrate and form a
mucilaginous liquid, which provides the way for the entry and union of antherozoid with the egg.
10.12.10 Sporophyte
It can be differentiated into foot, seta and capsule (Fig. 10.35A). The foot is bulbous, remains buried in
between the leaves at the apex of the female gametophore, and consists of thin-walled parenchymatous
cells. The seta is a very long and slender part of the sporogonium which pushes the capsule towards the
upper side, and its function is elongation.
194 � Bryophyta
Anatomically, the seta consists of an outermost superficial layer of thick-walled cells, followed by
one or a few layers of sclerenchymatous cells, which merge internally into parenchymatous, green and
thin-walled cells, having some intercellular spaces. Some very simple centrally-located cells form the
central strand of the seta.
The elongating seta enlarges at the base of the capsule in the form of a region called apophysis
(Fig. 10.35B). In between the apophysis and the sporogenous region (theca) is present a groove. The
stomata are present in the epidermis of the apophysis, specially in the region of the groove. Inner to the
epidermis are present some chlorophyll-containing cells in the region of apophysis.
In the theca region, the capsule wall consists of an outermost layer of epidermis, followed by a few
layers of chlorophyll-containing cells. Several air spaces traversed by chlorophyll-containing filaments
are present on the outer (outer lacuna) as well as the inner (inner lacuna) sides of the spore sac. The
spore sac, thus, separates the outer and inner lacunae. The inner air spaces thus remain in touch with
the centrally-located columella. When young, the archesporium is only unilayered but, in the mature
sporogonia, it is 4- to 6-layered. The archesporium or sporogenous cells develop into diploid spore
mother cells. Each spore mother cell divides meiotically into four haploid spores.
The operculum is a lid-like structure present at the top of the capsule (Fig.10.35 B,C). It contains
a conical beak or rostrum. Instead of annulus, a rim or diaphragm (Fig. 10.35C) is present. The
epiphragm, present at the base of the operculum, is a transverse band of thin-walled and compressed
cells with no intercellular spaces. The epiphragm actually closes the upper part of the capsule. The
peristome is present in the form of thick, fibrous, crescent-shaped cells near the epiphragm. In the
mature sporogonium, the peristome is represented by 32 or 64 peristomial teeth connecting the capsule
wall and the epiphragm. The peristomial teeth help in the dispersal of spores.
Fig. 10.35 A-C A, A female plant of Polytrichum commune bearing a mature sporogonium; B, LS of the capsule
of same; C, LS upper part of the capsule of P. commune showing details of theca and operculum

The spores are very large in number, but very small in size (0.005 to 0.01 mm in diameter), and
remain viable for a long time. According to Wigglesworth (1947), the yellow-coloured spores, at the
time of germination, become green due to the formation of chloroplast in P. commune. The endospore
ruptures the outer exospore and comes out in the form of one or more germ tubes. Repeated transverse
divisions in the germ tube result in a filamentous protonema. Some of these filaments penetrate into
the substratum and develop oblique walls, while other filaments grow towards upper sides. Some buds
develop from these upright filaments, which develop into young gametophores of Polytrichum.
1. What do you mean by the following terms?

(a) Bog (b) Peat
2. _____ are commonly known as bog mosses or peat mosses.
3. The most common bryophyte of bogs is_____.
4. Men on at least four points of differences between bog mosses and other mosses.
5. Sphagnales comprises of how many genera? Name one of them.
6. Is Sphagnum an annual moss or a perennial moss?
7. Why is Sphagnum also known as peat moss?
8. “Sphagnum is of great commercial importance”. Comment in about 100 words.
9. With the help of suitable illustra ons, describe the external features of mature gametophore
of Sphagnum.
10. Define the terms “innova on” in reference to Sphagnum.
11. Describe anatomy of leaf and stem of Sphagnum with the help of suitable illustra ons.
12. Explain following terms with reference to the anatomy of axis of Sphagnum:
(a) Hadrom, (b) Retort cells.
13. How does Sphagnum reproduce vegeta vely?
14. Describe in detail the sexual reproduc on in Sphagnum.
15. What do you mean by “primary archegonium” and “secondary archegonium” in Sphagnum?
16. Give an illustrated account of development of sporophyte in Sphagnum.
17. In Sphagnum, the archesporium is _____ in origin.
18. Explain the following terms with reference to the mature sporophyte of Sphagnum:
(a) Pseudopodium, (b) Vaginula
19. Explain explosive mechanism of dehiscence of capsule in Sphagnum.
20. Describe affini es of Sphagnum.
21. Men on at least six unique characteris cs of Sphagnum.
22. The common name “lantern mosses” is given to which category of mosses?
23. Write any five characteris c features of Andreaeales.
24. Describe the life history of Andreaea in about 500 words giving suitable illustra ons.
25. The name “granite moss” is given to which bryophyte?
26. Are structures like seta, operculum and peristome present or absent in the mature sporophyte
of Andreaea?
27. In mature sporophyte of Andreaea, the func on of seta is taken by a body of gametophy c
ssue known as _____
196 � Bryophyta
28. Write at least seven general characteris cs of Bryales.

29. The name “true mosses” is given to members of _____.
30. Give a brief illustrated account of the life history of Funaria in about 1000 words.
31. What is the most common species of Funaria, also available widely in India?
32. In “true mosses” (Bryales), is pseudopodium present?
33. Describe some methods of vegeta ve reproduc on in Funaria.
34. How can you differen ate between a gemma and a bulbil in Funaria?
35. Explain development of sporophyte in Funaria with the help of suitable diagrams.
36. Describe the fate of amphithecium and endothecium in the development of capsule of
Funaria.
37. Make well-labelled diagrams of the following parts of Funaria:
(a) LS capsule, (b) TS seta, (c) Peristomial teeth.
38. Write a botanical note on Buxbaumia in about 250 words.
39. Regarding size of Buxbaumia, is it a macroscopic or microscopic moss?
40. How many teeth does Tetraphis contain in its peristome?
41. What is the common name of Tetraphis?
42. Write a botanical note on Archidium in about 200 words only.
43. The name “large-spored-moss” is given to _____.
44. Describe external features and sporophyte of Pogonatum in about 300 words.
45. What is Polytrichum? Describe its life history in about 500 words. Substan ate your answer
with suitable diagrams.
46. In terms of rhizome anatomy of Polytrichum, explain the following terms:
(a) Hydroids, (b) Leptoids, (c) Hydrom, (d) Amylom
47. Describe mature sporophyte of Polytrichum with the help of suitable illustra ons.
11
Gametophyte of
Bryophytes
WHAT IS A GAMETOPHYTE? 11.1
A gametophyte is technically the haploid generation in an alternation of generations. The gametophyte
is the generation producing gametes. In bryophytes, the gametophyte is the main vegetative stage. In
angiosperms, however, the gametophyte is very small, contained in the ovule and pollen grain.
On the other hand, sporophyte is the diploid generation in an alternation of generations, and
it produces the spores. In bryophytes, the sporophyte grows directly from the archegonium of the
gametophyte, and it thus depends on the gametophyte for its nutrition.
EXTERNAL FORM OF MATURE GAMETOPHYTE 11.2

The mature gametophyte of bryophytes shows a wide range in its structure and size. There are bryophytes
in which the whole plant is very small and microscopic, i.e. Zoopsis argentea, while some bryophytes
are very large and attain a size as much as 50 cm in height (e.g. Dawsonia superba) and 50 to 70 cm
Fig. 11.1 A-D Gametophytes of some bryophytes. A, Zoopsis argentes; B, Buxbaumia aphylla;
C, Monoclea forsteri; D, Riccardia pinguis
198 � Bryophyta
in length (e.g. Fontinalis antipyretica). Buxbaumia aphylla (Fig. 11.1 B) attains a length of only a few
millimetres while Carrpos sphaerocarpos is spheroidal in shape and attains a diameter of 0.5 to 2 mm.
Monoclea foresteri attains a width of 5 cm. and a length of as much as 20 cm (Fig. 11.1 C).
Structurally, the gametophytic plant body of bryophytes may be simple thallus-like, i.e. thalloid, or
may bear leafy shoots, i.e. foliose.
THALLOID FORMS OF GAMETOPHYTE 11.3

A more or less undifferentiated plant body, without distinct roots, stem and leaves, is known as thallus.
In thalloid liverworts, the gametophyte is a flat, more or less
undifferentiated thallus. Thallus is found in members of
Marchantiales, many anacrogynous Jungermanniales and
Anthocerotopsida. It contains a distinct midrib, as in Riccia,
Marchantia and Dendroceros. Midrib is, however, absent in many
thalloid forms, such as Monoclea (Fig. 11.1C) and Riccardia
pinguis (Fig. 11.1D). Thalloid bryophytes are usually
dichotomously branched, e.g. Riccia and Marchantia. Closely
developing dichotomous branches form rosette-shaped structures,
as in Riccia.
Gametophytes of some bryophytes are neither typically thalloid
nor typically leafy forms. They appear to be intermediate forms
between thalloid and leafy forms, as in members of Metzgerineae
(acrogynous Jungermanniales), e.g. Treubia and Fossombronia. Fig. 11.2 Female gametophyte of
In Treubia insignis, the plant body has a thick, fleshy prostrate Fossombronia intes nealis
axis bearing two rows of well-developed fleshy leaves. In bearing archegonia
Fossombronia, the plant body is also leafy with two rows of
leaflike lobes borne on an axis (Fig. 11.2).
LEAFY FORMS OF GAMETOPHYTE 11.4

11.4.1 Leafy Forms of Hepa copsida
1. Sphaerocarpales The gametophytic plant body of Sphaerocarpos and Geothallus consists
of an axis containing two rows of alternately inserted lobe-like “leaves”. In Riella americana, the
gametophyte consists of a thick stem-like axis containing two rows of leaflike scales and a well-
developed undulate wing or two wings (Fig. 11.3A).
2. Calobryales In Calobryum blumei and Haplomitrium, the gametophytic plant body consists of
an underground rhizomatous region devoid of leaves and upright leafy shoots bearing three rows of flat
and unlobed leaves (Fig. 11.3B).
3. Takakiales In Takakia, the gametophytic plant body is made up of branched underground rhi-
zomatous portion whch lacks rhizoids, and erect axis with phyllids (Fig. 11.3C) or isophyllous leafy
shoots. Phyllids do not show any definite arrangement.
Gametophyte of Bryophytes � 199
Gametophyte Sporophytes
Leaves
Wing
Phyllid
Leafy
shoot
Axis
C
Axis B
A
D E G
F
Fig. 11.3 A-G, Gametophytes of some leafy forms of Hepa copsida. A, Riella americana; B, Calobryum
blumei; C, Takakia showing por on of axis with phyllids; D, Ventral view of Herber a adunca;
E-F, Dorsal and ventral view of Plagiochila asplenioides; G, Ventral view of Frullania armiliana
showing water sacs on lateral leaves
4. Jungermanniales In acrogynous Jungermanniales, the plant body usually consists of a pros-

trate to procumbent leafy axis bearing distichous leaves. In Herbertia (Fig. 11.3D) and a few more
genera, the leafy shoot shows radial symmetry. It bears an erect axis bearing three rows of radially ar-
ranged leaves. Usually, two rows of these leaves are laterally placed and a third row of smaller leaves
(amphigastria) are ventrally placed. In Plagiochila asplenioides (Fig. 11.3 E), the leaves are arranged
only in two rows. The smaller ventrally-placed leaves (amphigastria) are absent.
As far as the form or shape of the leaves in acrogynous Jungermanniales is concerned, they may
be entire (Porella), bilobed (Herbertia), trilobed (Bazzania), four-lobed (Lepidozia) or even highly
divided. In Frullania armiliana (Fig. 11.3 G), lateral leaves contain water sacs.
Two main types of branching in acrogynous Jungermanniales are terminal or end branching (as in
Frullania, Porella, Microlepidozia and Radula) and intercalary branching (as in Herbertia, Plagiochila,
Scapania and Cephaloziella).
11.4.2 Leafy Forms of Bryopsida

Gametophytic plant body of mosses is usually differentiated into rhizoids, stem and leaves. The
gametophores of different genera have different sizes, varying from very small and microscopic (e.g.
Ephemerum) to very large reaching up to 50 to 70 cm (e.g. Dawsonia).
The axis may be radial (e.g. Funaria) to dorsivental (e.g. Rhocopilum africans, Hylocomium
splendens).
200 � Bryophyta
The leaves are usually sessile and subulate to orbicular in outline in different species of mosses.
They are never lobed. When young, the leaves are spirally arranged in three vertical rows on the axis.
Each row of leaves corresponds to three cutting faces of the apical cell. In a few mosses, the leaves are
arranged only in two rows on opposite side of the axis (e.g. Distichium, Fissidens). The mature leaves
are spirally arranged with a divergence of 1/3 (Fontinalis antipyretica), 2/5 (Sphagnum), 3/8 (Funaria),
5/13 (Polytrichum), 4/11 (Dicranum scoparium) and 8/21 (Leskea).
Bryopsida members never show dichotomous type of branching. The branching is usually lateral,
but not axillary. On the basis of the two major types of branching, the mosses may be divided into
two groups, viz. acrocarpous and pleurocarpous. Acrocarpous mosses have an upright stem, with the
reproductive organs at the apex. On the other hand, pleurocarpous mosses have a stem with many
branches, spreading across the ground. The reproductive organs in pleurocarpous mosses are borne
on short side branches. A dendroid habit is resulted due to brancing in some mosses, e.g. Climacium
dendroides.
APPENDAGES FOUND ON THE GAMETOPHYTE 11.5

Rhizoids, scales, mucilage papillae (or hairs) and paraphyllia are some of the appendages found on the
gametophyte of bryophytes.
11.5.1 Rhizoids
A rhizoid is a thread-like cell which grows from the lower surface or base of a bryophyte. Bryophytes
lack roots. The rhizoids in bryophytes have the function of roots. Their combined functions include
anchorage and absorption. In a majority of Hepaticopsida and Anthocerotopsida, the rhizoids are
unicellular. But in Bryopsida or mosses, the rhizoids are multicellular, well-branched and contain oblique
septa. The unicellular rhizoids are of two types, smooth-walled and tuberculate, as in Marchantia, Riccia
(Fig. 7.2). In Anthoceros and some members of Jungermanniales and Sphaerocarpales, only smooth-
walled rhizoids and present. In some members (e.g. Riccia fluitans, Ricciocarpos natans, Takakia), the
rhizoids are absent. The bundles of rhizoids form anchoring discs in some members, e.g. Lejeunea. In
some bryophytes, the rhizoids develop in very large quantity and form a red or dark-brown felt-like
covering or tomentum over the stem, as in Dicranum scoparium and Mnium punctatum.
11.5.2 Scales
Multicellular, one-celled thick, purple-coloured structures found on the ventral surface of Marchantiales
(e.g. Riccia, Marchantia; Fig. 7.17) are known as scales. Scales are absent in other members of
Hepaticopsida, Anthocerotopsida and Bryopsida.
The scales usually protect the growing point of the thallus. They keep the growing points moist and
thus protect them from the loss of water. The usual purplish colour of the scales is due to the presence
of anthocyanin pigment. Small-sized chloroplasts are also present in the scales of many members,
including Asterella reticulata, Athalamia pusilla, Cryptomitrium himalayensis and Stephensoniella
brevipedunculata. Due to the presence of such chloroplasts, the scales are also assimilatory in function
in these bryophytes. Scales are either simple and ligule-like, as in Riccia. But in Marchantia, scales are
of two types, i.e. ligulate and appendiculate. They are either arranged in one row (e.g. Riccia), in two
rows on either side of the midrib (e.g. Oxymitra), or in two to four rows on either side of the midrib (e.g.
Marchantia). In some bryophytes, the scales are irregularly distributed, as in Athalamia and Corsinia.
11.5.3 Mucilaginous Hairs or Mucilage Papillae

Mucilaginous hairs or papillae secrete mucilage, which protect the growing points against drought.
They are found in some Hepaticopsida, e.g. Sphaerocarpos, Blasia, Metzgeria, Marchantia, Lunularia,
etc. Mucilage hairs are usually absent in Anthocerotopsida. In Sphaerocarpos, the mucilage hairs
develop on the ventral surface of the thallus near the growing point. In Blasia, they are found on both
the surfaces of the thallus. Mucilage hairs may be unicellular (as in some species of Sphaerocarpos) or
multicellular (as in Calobryum and Diplophyllum). In Takakia, the mucilage hairs are of two types: (i)
filamentous closed type, and (ii) beaked open type.
11.5.4 Paraphyllia
Paraphyllia are the delicate minute appendages of variable forms, found on the stem and sometimes on
the leaf bases of some Hepaticopsida (e.g. Trichocolea paraphyllina) and Bryopsida (e.g. Plagiothecium
deplanatum). These are made of chlorophyll-containing tissue, and found intermixed with normal
leaves. In Bryopsida, the paraphyllia are also found in some species of Ectropothecium, Helodium,
Hylocomium and Thuidium. Paraphyllia have the ability to absorb and retain water like a sponge, and
also help in external capillary condition of water. Because of the presence of chlorophyll, they may also
be helpful in photosynthetic activity of the plant.
INTERNAL FORM OF MATURE GAMETOPHYTE 11.6

11.6.1 Anatomy of Mature Gametophyte
Anatomy of thalloid forms of gametophyte of bryophytes is highly variable in different members.
In very simple forms (e.g. Anthocerotales, thalloid Jungermanniales and some Marchantiales), the
thallus is made up of one to several layers of uniform, undifferentiated cells, with no differentiation of
tissues. In Pellia, the cells of upper layer of the midrib and wings usually contain chloroplasts, which
are usually absent in other cells of the thallus (Fig. 11.4).
Fig. 11.4 Transverse sec on of the midrib region of the thallus of Pellia fabbroniana
202 � Bryophyta
In some thalloid Jungermanniales (e.g. Hymenophytum, Makednothallus, Pallavicinia and

Podomitrium), the midrib is transversed by a centrally-located strand made up of narrow, elongate,
thick-walled cells with pointed ends. These cells are densely pitted and usually lack protoplasmic
content. The centrally-located strand of these genera functions as a water-conducting system. Smith
(1964) compared this strand with the xylem of vascular plants. It, however, lacks tracheids and vessels,
characteristic of xylem.
More anatomical differentiation of tissues is observed in a majority of Marchantiales. There is a
clear anatomical differentiation of dorsal and ventral regions. In ventral region, the thallus contains
large, parenchymatous cells containing starch. It functions as the storage region of the food reserves.
Mucilaginous cells and oil bodies may also be present in the ventral region in genera like Marchantia
and Targionia. The dorsal region is mainly photosynthetic in function because it consists of chlorophyll-
containing cels. A network of large air chambers is present in the entire thallus of aquatic species such
as Riccia fluitans (Fig. 7.1C) and Ricciocarpos natans.
In Conocephalum, Lunularia, Marchantia and Targionia, the dorsal region of the thallus contains
a single horizontal layer of air chambers, just below the upper epidermis. Air chambers are separated
by vertical partitions in these genera. In Asterella and Reboulia, the air chambers are in several rows,
and they are separated by a single layer of chlorophyll-containing cells. Secondary partitions may also
be present in these air chambers. The air chambers, when present in several rows, are always empty,
and do not contain chlorophyll-containing cells.
Air chambers open on the dorsal surface of the thallus, and these openings are called air pores.
In Riccia, the air pores are simple air spaces bounded by 4 to 6 or more cells. In many other thalloid
bryophytes, the air pores may be simple or barrel-shaped or compound (e.g. Marchantia). In simple air
pores, the pore is either surrounded (i) by a single ring of 5 to 8 cells and such pores appear star-shaped
in surface view, as in Peltolepis; or (ii) by one or two rings of thin-walled cells, as in Corsinia; or (iii)
by 3 to 8 concentric rings of cells, of which each ring is made up of 6 to 8 cells, as in Plagiochasma,
Reboulia, etc. The compound air pores of Marchantia contain 4 to 5 superimposed concentric rings of
cells, of which each ring consists of 4 to 8 cells.
11.6 2 Anatomy of Leafy Forms of Gametophyte
1. Hepa copsida
Leafy forms of the gametophytes of Hepaticopsida consists of stem and leaves.
(a) Stem A well-differentiated conducting strand is absent in the leafy Hepaticopsida. They are
able to absorb water through any of the external surface of the stem, and such members are called
ectohydric.
In leafy forms of Hepaticopsida, the stem is usually composed of uniform cells with no differentiation
of tissues. The size and thickness of the walls of superficial or cortical cells and internal or medullary
cells, however, differ in different members, as exemplified by the following:
(i) In Bazzania and Omphalanthus, the cells of the cortical layer are smaller than that of the
medullary cells. Almost all cells of the stem have more or less thickened walls.
(ii) In Frullania, Plagiochila and Scapania, the cortical cells are smaller and thick-walled while
the medullary cells are usually larger and thin-walled.
(iii) The stem of Acromastigum contains 7 rows of large cortical cells.
(iv) The stem of Zoopsis contains 4 to 6 rows of cortical cells.

(v) The stem of Lejeunea contains 7 rows of cortical cells and as many as 10 to 20 rows of
medullary cells.
(vi) The stems of Drepanolejeunea and Leptolejeunea contain 7 rows of cortical cells and only 3
rows of medullary cells.
(vii) In Cololejeunea, the stem contains 5 rows of cortial cells and only 1 row of medullary cells.
(viii) In Calobryum, the stem contains an outer zone of cells containing oil droplets and pale green
plastids and a central zone of colourless elongated cells. A thin cuticle is also present on the
outermost side.
(ix) In Takakia, the cortical cells are usually thick-walled while the medullary cells are thin-
walled.
(b) Leaf Usually, the leaves are one layer of cells in thickness, as in Calobryum and Haplomitrium.
The leaves of the basal part of the plant in these members are, however, two to four cells in thickness.
The cells of the leaves are parenchymatous. Leaves of leafy Hepaticopsida usually lack a midrib. In
some genera, the leaves are pluristratose as in Chondrophyllum cucculatum.
The shape of the cells of leaves is variable. It may be round (e.g. Bazzania trilobata), rectangular
(e.g. Blepharostoma trichophyllum) or even polygonal (e.g. Calypogeia trichomanis). Numerous
chloroplasts are present in each cell of the leaf.
2. Bryopsida
(a) Stem In Bryopsida (mosses), the stem is generally differentiated into an epidermis, cortex and
central conducting strand. In several mosses, however, there is no well-differentiated conducting
strand, e.g. Drummondia, Hedwigia, Helodium, Neckera and Ulota. Anatomically, the stem shows
several variations, a few of which may be exemplified as under:
(i) In Sphagnum, the central cylinder is surrounded by cortex. The conducting strand is absent.
(ii) In Andreaea, the stem contains uniformly thick-walled cells without any differentiation of
cortex and central conducting strand.
(iii) In Polytrichum, the rhizome contains an endodermis-like layer and a pericycle. Its central
cylinder shows differentiation of tissues.
(iv) In Dawsonia also, there is a differentiation of tissues in the central cylinder, like Polytrichum.
Both these genera also contain leaf traces which are loosely connected with the central
cylinder.
(v) In Bryum and Mnium, a distinct conducting strand is present. Water in these mosses is
absorbed by the rhizoids and conducted up to leaves through the stem. Such mosses are called
endohydric.
(vi) In Funaria and several other mosses, the cortex of the stem contains leaf traces. Such traces
end bindly in the cortex without reaching up to the central cylinder.
(b) Leaf The leaves of most of the mosses possess a midrib, as in many species of Tetraphis, Fissidens,
Phascum, Lepidopilum and Callicostella. In several mosses, however, the midrib is absent, as in many
species of Sphagnum, Andreaea, Ephemerum, Fontinalis, Hedwigia and Nanomitrium. The midrib,
when present, is single in each leaf. But in Lepidopilum and Callicostella, each leaf possesses two
midribs. In Fissidens bryoides, the midrib may extend up to the tip of the leaf.
204 � Bryophyta
The leaves of mosses is usually one-celled thick except the midrib region. The cells of this single-
layered wing or laminate portion contain chloroplasts. In Polytrichum, the leaves show highest grade
of development. In a majority of mosses, the apical cell of the leaf is wedge-shaped, and usually it cuts
segments from two sides.
1. Give an illustrated account of the external form of mature gametophyte of bryophytes.

2. Write an essay on gametophyte of bryophytes in about 1000 words.
3. In an alterna on of genera ons, a gametophyte is technically a _____ genera on.
4. In bryophytes, the _____ is the main vegeta ve stage.
5. _____ is the diploid genera on in an alterna on of genera ons, and it produces the _____.
6. Name a very small-sized, microscopic bryophyte.
7. The gametophy c plant body of bryophytes may be thalloid or _____.
8. Describe some leafy forms of the gametophyte of Hepa copsida giving suitable illustra ons.
9. Describe the appendages found on the gametophyte of bryophytes.
10. Write short notes on:
(a) Paraphyllia, (b) Mucilage papillae, (c) Scales.
11. Describe the anatomy of mature gametophyte of bryophytes.
12. How can you differen ate between ectohydric and endohydric mosses?
12
Sporophyte of
Bryophytes
WHAT IS A SPOROPHYTE AND A SPORE? 12.1
From the technical viewpoint, the sporophyte is the diploid generation in an alternation of generations.
The sporophyte is the generation producing spores, which are haploid. In angiosperms, gymnosperms
and pteridophytes, the sporophyte is the main vegetative stage. In bryophytes, on the other hand,
the sporophyte grows directly from the archegonium of the gametophyte, and thus depends on the
gametophyte for its nutrition.
Spore is a small round cell from which a whole new plant is produced. In bryophytes, the spores
are haploid and are produced by the sporophyte. They are produced as a result of meiosis in the spore
mother cells
In bryophytes, the sporophyte is usually made up of three parts, viz. foot, seta and capsule. Foot
is the basal part which remains embedded within the gametophyte. It is basically an absorbing and
anchoring structure. Seta is an elongated, stalk-like structure and bears the third most important part of
the sporophyte, the capsule. Foot and seta are sterile structures while the capsule is the fertile structure
of the sporophyte which produces the spores.
HOW IS THE SPOROPHYTE DIFFERENT

FROM THE SPOROGONIUM? 12.2
Capsule is the spore-bearing structure of the sporophyte in bryophytes, and it has also been described as
sporogonium. The terms like “sporogonium” and “sporangium” are used as synonyms of the capsule.
Since, the sporangium is the spore-bearing structure in pteridophytes, its use in bryophytes should be
abandoned or stopped. The term “sporogonium” is usually used as an alternative for a sporophyte in
bryophytes. As mentioned in the Chambers Biology Dictionary, the term “sporogonium” is the “same
as the sporophyte in bryophytes”. In Longman’s Illustrated Dictionary of Botany, sporogonium is “the
sporophyte of a moss and a liverwort consisting of a foot, seta and capsule.”
206 � Bryophyta
BRYOPHYTIC SPOROPHYTE: A BODY DEPENDENT ON

GAMETOPHYTE 12.3
In bryophytes, the sporophyte is always dependent on the gametophyte, and due to this some bryologists
even describe it as “a parasite on the gametophyte”. However, since some cells of the wall of the capsule
contain chloroplasts, the sporophyte is not fully dependent for nourishment on the gametophyte, and
due to this it should not be described as a parasite on the gametophyte. In some bryophytes, the cells
of the seta also contain chloroplasts. A few cells of the foot also contain some chloroplasts in a few
bryophytes, such as Sphaerocarpos. In comparison to mosses, the chlorophyll content in the cells
of seta and capsule is very low in liverworts. Due to the presence of chlorophyll in some cells, the
sporophyte is not fully dependent on the gametophyte. It is partially dependent.
In Anthoceros, the sporophyte is highly organised, green throughout its life and even some of its
epidermal cells contain functional stomata, helpful even in gaseous exchange. A well-organised region
of mechanical support, also helpful in conduction, is present in the form of columella in the sporophyte
of Anthoceros. Some intercalary meristematic cells are also present in the basal part of the sporophyte.
All these make the Anthoceros sporophyte a highly organised and highly evolved body.
STRUCTURE OF SPOROPHYTE 12.4

As mentioned earlier, the sporophyte in most liverworts (e.g. Marchantia; Fig. 12.1A) and mosses is
made up of foot, seta and capsule. A brief discussion of all these parts is mentioned below.
Fig. 12.1 A, A mature sporophyte of Marchan a showing foot, seta and capsule;
B, A leafy shoot with sporophyte and enlarged marsupium
Sporophyte of Bryophytes � 207
12.4.1 Foot
The base of the sporophyte of a bryophyte, which is the part that attaches it to the gametophyte, is
known as foot. It functions for absorption and anchorage. Its size and shape are variable in different
bryophytes. In some bryophytes, the foot is absent, e.g. Riccia. In most bryophytes, the foot is globose
(e.g. Conocephalum, Corsinia, Marchantia, Anthoceros) to anchor-shaped (e.g. Pellia, Porella). In a
majority of thalloid liverworts, the foot is large and massive, but in a few liverworts it is very small
(e.g. Trichocolea). The foot is nearly spherical and bulbous in a few mosses (e.g. Micromitrium). In
Schistochila, a foliose liverwort, some rhizoid-like extensions develop from the foot. Marsupium, a
tuber-like outgrowth of the foot, is seen in Calypogeia (Fig. 12.1 B).
12.4.2 Seta
Seta is the stalk of the sporophyte of a bryophyte. When young, the seta is made up of elongate cells
meant for conduction and support. Seta is absent in some bryophytes, e.g. Riccia, Anthoceros and
Notothylas.
The seta usually elongates by elongation of its cells and not by division of its cells. Usually, the
elongation of the seta is as fast as 1 mm per hour, when young. It attains a length of over 5 cm
or more in some liverworts, e.g. Pellia, Monoclea, etc. In Marchantia (Fig. 12.1A), Corsinia and
Targionia, the seta is not a very long structure. In Pohlia natans, seta reaches up to 7 cm in length,
and in Drepanocladus fluitans, seta may attain a length up to 10 cm or even more. In genera such as
Phascum and Ephemerum seta is a very minute body.
Seta is long but a delicate and ephemeral structure, made up of thin-walled cells. Transverse section
of the seta of Jubula reveals that in circumference, it is only made up of a few cells (Fig. 12.2A). But
it is made up of many cells in Mylia (Fig. 12.2 B). In Caphaloziella, seta consists of four small central
cells surrounded by four larger cells (Fig. 12.2 C).
Fig. 12.2 Transverse sec ons of the seta of Jubula (A), Mylia (B) and Cephaloziella (C)
Some of the striking differences between seta of liverworts and mosses are listed in Table 12.1.
In several species of Brachythecium and Dicranella, the seta contains some specialised papillae-like
structures. In Brachythecium rutabulum, papillae on the seta are quite large and can be seen even with
the naked eye.
208 � Bryophyta
Table 12.1 Differences between the seta of liverworts and mosses
S NO. LIVERWORTS MOSSES

1. Seta is quite an elongate body much before the Seta is a short structure prior to the maturity of
maturity of capsule. capsule.
2. Seta is not highly specialised for conduction. Seta is a highly specialised structure for conduc-
tion.
3. It does not provide that much support to the capsule It provides more support to the capsule than in
as in mosses. liverworts.
4. It is more significant for dehiscence in liverworts. It is not so significant for dehiscence in mosses as
it is in liverworts.
12.4.3 Capsule
Capsule is the spore-producing organ of the sporophyte of a bryophyte, borne at the top of the seta. In
liverworts, it varies in its form in different members. It may be cylindrical, ovoid, subspherical or even
spherical. In Riccia (Fig. 7.10 J), Sphaerocarpos (Fig. 5.2 M), Frullania, Porella (Fig. 4.7), Pellia (Fig.
4.16 B), Fossombronia, Lejeunea and some other bryophytes, the capsule is nearly spherical in shape.
Capsules are almost ovoid or ovoid-cylindrical in Blasia and Riccardia while they are elongated in
Haplomitrium and Monoclea.
As far as the size is concerned, the capsules of mosses are usually larger than the capsules of
liverworts. In Marchantiales, they attain a diameter of 1 to 1.25 mm but in Jungermanniales, the
capsules are comparatively narrower and reach up to 0.6 to 1 mm in diameter. In Riccardia and Pellia,
the capsule reaches up to 1.5 mm or more in diameter.
In mosses, the capsule is usually cylindrical and erect (Funaria hygrometrica), subglobose (e.g.
Bartramia) or pyriform and pendulous (Bryum, Pohlia).
DEVELOPMENT AND DIFFERENTIATION OF

DIFFERENT ORGANS OF SPOROPHYTE 12.5
Development and differentiation of different organs of the sporophyte in bryophytes may be studied
in the form of five different stages, viz. embryogeny stage, protected stage, green stage, differentiation
stage and, dehiscence and dispersal stage.
12.5.1 Embryogeny Stage

A plant at an early stage of development is known as embryo, and the processes leading to the formation
of the embryo is known as embryogeny. So, early developments of the sporophyte of a bryophyte are
included under the embryogeny stage.
Two main findings of the embryogeny stage in liverworts are (i) early embryo passed through
the formation of an 8-celled octant stage (Marchantiales) or linear stage (Jungermanniales), and (ii)
origin of sporogenous tissue. The young multicellular embryo divides periclinally to form the outer
amphithecium and the inner endothecium, either of which form the sporogenous tsissue in different
members. Endothecium is, however, responsible for the formation of sporogenous tissue in most
liverworts.
In mosses, on the other hand, the early embryogeny (Fig. 12.3 A-H) is more uniform. Polarity is
established in the initial stages, and elongation of the young sporophyte takes place by the activity of
an apical cell. In the lower part of the young sporophyte, a second apical cell starts functioning. Soon,
the ovoid embryo of the moss transforms into an ellipsoidal body and finally a narrow cylindrical body
tapering at both ends is resulted. The central endothecium is soon demarcated from the peripheral
amphithecium in this young multicellular cylindrical embryo. The archesporium originates from
the endothecium. The external wall layers and central columella are differentiated when the young
embryo is about 20 to 24 cells thick. Various mosses require different periods for the differentiation
and development of various parts (i.e. foot, seta and capsule) of the sporophyte. For example, the
processes from the time of fertilization up to the discharge of spores are completed within 2–3 weeks
in Phascum cuspidatum, but in Polytrichum the period required for completion of all these processes
is over one year.
Fig. 12.3 A-H Diagramma c representa on of the early stages of embryogeny in a moss capsule.
A, Two-celled stage; B, Quadrant stage; C-D, Differen aion of amphithecium and
endothecium; E-H, Differen a on of wall layers, sporogenous layer and columella
12.5.2 Protected Stage

Calyptra and positioning of foot are the two major ways which help in the protection of young
developing sporophyte. Besides calyptra and foot, some additional protective structures are also
present in some bryophytes. Involucre, green curtain of tissues and perianth are some such additional
protective structures. Well-developed pear-shaped involucres are present as a protective body in
Sphaerocarpos. Some green curtain of tissues develop amongst the archegonia in Corsinia. Some
bryologists consider these tissues as forerunners of carpocephalum. Infoldings of scale-like structures
develop as a protective tissue in the sporophytes of Targionia. A green involucre is present in Pellia. In
Marchantia, the protective structures of sporophyte include calyptra, perianth and involucre. A Chinese-
lantern-type additional protective covering is present in the sporophyte of Fimbriaria. Riccardia lacks
any additional protective structure.
210 � Bryophyta
1. Calyptra
The calyptra is actually a hood of tissue produced from the wall of the archegonium, especially in
mosses. It is also formed in liverworts. Calyptra protects the young sporophyte. The size and shape of
the calyptra affect the shape and orientation of the capsule in mosses. The calyptra is highly variable
in different bryophytes in its extent of covering the capsule. In bryophytes, in general, and mosses in
particular, the shape of the calyptra serves as a useful tool of taxonomic importance. In Eucalypta,
the calyptra is an elongate cone-like structure which covers the capsule all over. In a majority of other
bryophytes, the calyptra is a small cap-like covering surrounding the basal part of the developing capsule.
Due to very fine hairy calyptra in Polytrichum, the name hair-cap-moss is given to this genus.
2. Foot
The foot of the sporophyte of bryophytes remains housed in the green gametophytic tissue. It provides
adequate nutrition to the sporophyte. In some leafy liverworts, the sporophyte is housed in a special
pouch-like structure of gametophytic origin, known as marsupium. It is a multilayered pouch-like
or tube-like structure made up of gametophytic tissue. In Geobelobryum, the marsupium is a well-
developed, elongated, tube-like structure bearing rhizoids, which are sometimes seen even buried into
the substratum like that of a root of higher plants.
In thalloid liverworts, the foot of the sporophyte is a globose mass of undifferentiated cells. In the
foot of mosses, some differentiation may be observed in the form of outer haustorial cells, intermediate
unspecialised cells and central cells. The central cells function like that of conducting cells. Intense
enzymatic activity can be observed in the cells of the outer region of the foot of mosses. Chauhan
(1988) reported cytochemical reactions for respiratory enzymes and phosphotases in the cells of the
foot region of Physcomitrium cyathicarpum. The haustorial cells of the foot of the sporophyte function
as the organs of absorption and transfer of nutrients, and due to these, they are named transfer cells. In
Physcomitrium cyathicarpum and Dendroceros, the peripheral cells of the foot show some infoldings
or invaginations, which form wall labyrinths. The characteristic feature of transfer cells is the presence
of these wall labyrinths in this genus. The transfer cells of the foot, therefore, form the junction between
sporophyte and gametophyte. Besides mosses and liverworts, transfer cells have also been reported in
hornworts (e.g. Anthoceros punctatus) by Ligrone and Gamberdella (1988). Transfer cells are absent
in the foot region of Pellia and Sphagnum.
12.5.3 Green Stage

Photosynthetic efficiency of the sporophyte represents its green stage. It is an important stage because
it provides some independence to the sporophyte from the gametophyte. In almost all liverworts,
the young sporophyte remains buried in the gametophytic tissue. In developing and nearly mature
sporophytes, the capsule is pushed out of the gametophytic tissue due to the meristematic activity of
seta and absorbing nature of foot. However, due to low chlorophyll content in seta and capsule, the
sporophyte depends on the gametophyte for its requirements of inorganic and organic contents. In
mosses, on the other hand, more amount of chlorophyll is present in different tissues and parts of the
sporophyte. The sporophyte is, therefore, more photosynthetically efficient than liverworts. The long
seta of the sporophyte of mosses is quite green, especially when young, and hence more efficient in terms
of its photosynthetic activity. The seta, however, is more specialised for support and conduction.
In mosses (e.g. Bartramia, Bryum, Funaria), a well-developed green tissue is present in the wall
of the capsule of the sporophyte. More photosynthetic activity is possible in the sporophytes of these
mosses. In Torula and some more mosses, the green tissue in the capsule region is less extensive, hence
there is less amount of photosynthetic activity. In Splachnum ampullaceum and many other species of
this genus, the apophysis region is most extensive amongst mosses.
Stomata are not found on the capsules of liverworts. In most mosses, they are present on the capsule,
specially in the apophysis region. In Pleuridium, the number of stomata are only 3-5 on a capsule. But
in some mosses (e.g. Philonotis), each capsule has as many as 200 or more stomata. Some mosses have
no stomata on their capsules, e.g. Fontinalis and Atrichum.
12.5.4 Differen a on Stage

Differentiation stage of the sporophyte is the demarcation of tissues which results into the formation of
sporogenous cells and spores in the sporophyte. The entire sporogenous tissue is endothecial in origin
in Marchantiales (e.g. Riccia) and results in the formation of spores. In Jungermanniales, a part of
the sporogenous tissue remains sterilised and forms the elaters. And, as we proceed further in mosses
the capsule elaborates and only very little of its tissue is meant for production of spores. Moreover, the
sterile structures like elaters are also not formed in capsules of mosses. A sterile region in the capsule
of Funaria and other mosses is present in the form of columella, which provides mechanical support
to the capsule and also helps in conduction. Columella is absent in liverworts. In Anthoceros and other
hornworts, a well-developed columella is also present.
The sporogenous tissue develops into spores, and the number of spores per capsule per plant have
been estimated in some bryophytic genera by bryologists. Some such findings are listed in Table 12.2.
Table 12.2 Approximate number of spores per plant in some bryophytes
S NO. GENUS NUMBER OF SPORES/SPORE TETRADS PER PLANT PER CAPSULE (APPROX.)
1. Sphaerocarpos 200 spore tetrads per capsule
2. Pellia 4500 spores per capsule
3. Lophocolea cuspidata 24,000 spores per capsule
4. Diplophyllum albicans 40,00,00 spores per capsule
5. Eurhynchium 700,000 spores per capsule
6. Scapania undulata 10,00,000 spores per capsule
7. Marchantia polymorpha 7000,000 spores per plant
12.5.5 Dehiscence and Dispersal Stage

Structure of the capsule wall is of utmost importance in the dehiscence of sporophyte in bryophytes.
Marchantiales possess a unistratose capsule wall whereas in Jungermanniales the capsule wall is
multistratose. The unistratose capsule wall of Marchantiales sometimes has transverse thickenings
(e.g. Peltolepis quadrata, Fig. 12.4A). But in Plagiochasma intermedium (Fig. 12.4B), the cells of the
jacket are thin-walled and do not bear any ornamentation. In Asterella, the capsule jacket is without an
apparent thickening (Fig. 12.4 C). But in Conocephalum, the entire transverse wall is thickened (Fig.
12.4 D). The wall of Norwallia shows ornamentation in the longitudinal wall (Fig. 12.4 E) while the
longitudinal wall shows thickenings (Fig. 12.4F). The epidermal cells shows ornamentation in case of
Frullania (Fig. 12.4 G) but in Radula (Fig. 12.4H), the epidermal cells of the wall of the capsule show
212 � Bryophyta
Fig 12.4 A-H Wall of capsule of Peltolepis quadrata (A) showing transverse thickenings; capsule walls of
Plagiochasma (B), Asterella (C), Conocephalum (D), Norwallia (E), Arnellia (F), Frullania (G),
and Radula (H)
thickenings. It has been observed in different members that the jacket of the capsule ruptures along
four longitudinal lines resulting in the formation of four valves or flaps. Intact capsules of Cephalozia,
before (Fig 12.5A) and after dispersal Fig. 12.5B) and in Frullania before (Fig. 12.5 C,D) and after
dispersal (Fig. 12.5 E) are shown in Fig. 12.5.
Fig. 12.5 A-B, Intact capsules before and a er dispersal in Cephalozia; C-E, Intact capsules
before and a er dispersal in Frullania
In mosses, the jacket of the sporophyte is multistratose. The stomata are also present in its epidermal
wall in most mosses. The stomata when present, are either exposed (e.g. Funaria) or immersed (e.g.
Orthotrichum). Operculum, a cap-like structure, gets differentiated in the apical region of the jacket of
the capsule. The operculum ruptures and this triggers the dehiscence of the capsule. The operculum is
released by the annulus, which is made up of a ring of enlarged elastic cells. The shape and structure of
the operculum help in the dehiscence and dispersal of spores.
1. Categories of Dehiscence of Capsule and Dispersal of Spores
Two categories of dehiscence of capsule and dispersal of spores may be as under:
(a) Passive Dehiscence and Passive Dispersal This takes place in those bryophytic genera (e.g. Corsinia,
Riccia) in which the seta is absent in the sporophyte and dehiscence takes place by disintegration of
jacket of the sporogonium. Because of the absence of elaters or any specialised structures, the dispersal
is also passive.
(b) Ac ve Dehiscence and Ac ve or Passive Dispersal This takes place in those genera in which the
sporogonium wall ruptures along four lines of dehiscence and elaters are present. The elaters help in
the dispersal of spores.
2. Dehiscence and Dispersal in Liverworts

Dehiscence of capsule and dispersal of spores in liverworts mainly take place by three different
mechanisms, described below:
(a) hygroscopic mechanism (e.g. Pellia, Marchantia, etc.),
(b) water-rupture mechanism (e.g. most of the foliose liverworts), and
(c) spiral-ring mechanism (e.g. Frullania).
Spores are dispersed slowly in hygroscopic mechanism but very fast and violently in water-rupture
mechanism and spiral-ring mechanism.
In members showing hygroscopic mechanism, the spiral bands of elaters are weaker than those
showing the other two mechanisms. On the other hand, in members showing water-rupture mechanism
and spiral-ring mechanism, spiral bands of their elaters show great strength of bispiral thickenings, as
in Cephalozia and Lophocolea. In dry conditions, the coils of bispiral thickenings contract sharply,
resulting in extreme tension. This loosens the coils abruptly, untwists the elaters instantaneously and
results in dehiscence and violent dispersal of spores.
In the liverworts showing hygroscopic mechanism resulting into slow dispersal of spores, the elaters
bear weaker spiral bands. The spirals of bands of these elaters, of course, contract but this contraction
is not sufficient to induce water rupture. The elaters simply twist in the available atmospheric humidity,
and there is seen only gradual dispersal of spores.
The spiral-ring mechanism is characteristic of Frullaniaceae and Lejeuneaceae, and this is linked
with unique internal organisation of the capsule. A series of elaters are present from roof to floor in
the globose capsule of these members. Dehiscence mechanism in these members is also unique. It
takes place by four valves of its jacket, which curve outwards rapidly, resulting into the discharge of
spores.
3. Dehiscence and Dispersal in Mosses
Various types of dehiscence of capsules and dispersal of spores are observed in mosses. In Andreaea
(Fig. 12.6A,B), the dehiscence of capsule is of great taxonomic significance. The jacket of the capsule
214 � Bryophyta
of this moss is thick-walled and possesses four or more lines of weakness which extend from the base
towards the apex. The mature sporogonium starts drying out, shrinks, and the shrinkage leads to lines
of weakness and finally yields into the dispersal of spores.
Fig. 12.6 Andreaea rupestris showing en re sporogonium (A) and LS (B) of the same
In Ephemerum and some species of Physcomitrium, the capsules are closed or cleistocarpous. They
open in an irregular manner to disperse the spores. Peristome and a detachable operculum or lid are
absent in such mosses. Seta is also ill-developed or even absent in these mosses. Only a few leaves
present in the gametophyte of these mosses surround the sporophyte. In dry conditions, the entire plants
of these mosses can be blown away by winds and are also transported far by humans and animals.
In gymnostomous mosses (e.g. Pottia truncata), the operculum may or may not be present in the
capsules. They also lack peristome. Due to these characteristics, these mosses have no gradual or
regulated mechanism of dispersal of spores. Dehiscence of the capsule takes place by blowing off of
the upper part of the capsule. The spores are dispersed by gravitational force because the capsule mouth
is directed downwards in these mosses
According to Edwards (1980), weather plays an important role in the dehiscence of the capsule and
dispersal of spores in aquatic mosses like Scouleria and Wardia. It is so because their capsules are
not directed downwards and they also lack teeth. In dry weather, their lid opens but in wet weather, it
remains intact, and spores are exposed due to wind.
Peristome is the major part to regulate the spore dispersal in the moss capsule, e.g. Funaria
hygrometrica. Two well-defined rings of peristome are present in this moss. In some other species
(e.g. Funaria fascicularis), however, the peristome is either missing or rudimentary. Similarly, two
rings of peristome are present in Encalypta streptocarpa but only one ring of peristome is present in E.
rhabdocarpa. Peristomate mosses thus fall in two categories, viz. haplolepideae (having single ring
of peristome), and diplolepideae (having two rings of peristome). Usually, a single ring of peristome
has 16 teeth. They form a close-fitting circle at the base and their apical parts taper to a point. Each
tooth of the peristome is a barred structure, and the tooth bars are actually remnants of the cell wall.
Thickenings, which are characteristic of outer teeth, are usually absent in the inner teeth. Usually,
16 teeth of the inner ring alternate with the 16 teeth of the outer ring. 16 thread-like cilia, in groups of 3
or 4, are also usually present on the same radii as that of the outer teeth. Dispersal of spores is actually
regulated by the peristome.
Regarding variations in peristomial teeth (Fig. 12.7 A-F), solid type of 4 erect peristome teeth are
present in Tetraphis (Fig. 12.7A), a ring of spirally twisted filliform teeth are present in Barbula (Fig.
12.7B), a ring of 16 barred slow-moving teeth are present in Dicranella (Fig. 12.7C) while double
peristome quite active in spore dispersal, are present in Hypnum (Fig. 12.7D) and Bryum (Fig. 12.7 E).
Peristome shows teeth curving over epiphragm in Atrichum (Fig. 12.7F).
Fig. 12.7 A-F Showing varia ons in peristomial teeth in Tetraphis (A), Barbula (B), Dicranella
(C), Hypnum (D), Bryum (E) and Atrichum (F)
216 � Bryophyta
The entire process of spore dispersal in mosses can be grouped in three broad categories, as under:
(a) Primi ve Type In mosses like Barbula (Fig. 12.7B) and Tortula, teeth movements have very little
or no active role in spore dispersal, and this is called primitive type. The peristome in such mosses
serves only as a hygroscopic lid.
(b) Intermediate Type In Dicranella (Fig. 12.7(C) and some other mosses, the peristome has 16
forked and freely moving teeth. In these mosses, the spores accumulated under the capsule mouth are
caught in between slowly moving teeth and get dispersed.
(c) Advanced Type Gradual discharge of spores is seen in the advanced type of mosses. Teeth play
a more active role in this type of dispersal. A majority of the mosses, which fall under this category,
have two rings of peristomial teeth. Advanced type is exemplified by mosses such as Brachythecium,
Bryum (Fig. 12.7E), Hypnum (Fig. 12.7D) and Mnium. The spores are dispersed in dry conditions in
these mosses by a process in which inner peristomial teeth stand up as a central cone and the tips of
teeth of outer ring are inserted into the gaps present in between different inner structures. In moist
conditions, the process of gradual discharge of spores depends on the ability of peristomial teeth to
close the capsule mouth.
In Atrichum (Fig. 12.7(F), Oligotrichum and Polytrichum, the peristome is of complex type, and
dispersal of spores takes place by censor mechanism. As many as 32 or 64 peristomial teeth of
different structures, are present in these mosses. Each tooth consists of fibre-like cells of several
layers of thickness, and this represents a solid construction. All these teeth unite at their tips to form
a membranous structure called epiphragm. Dispersal of spores takes place through very fine holes
present between the successive teeth, and this type of dispersal of spores is called censor mechanism.
An unusual type of spore dispersal is observed in Tetraphis (fig. 12.7A). Its peristome contains four
large teeth of solid construction, but it lacks epiphragm.
Sphagnum (Fig. 12.8A-D) shows air-gun mechanism of dehiscence of capsule and dispersal
of spores. The mature sporogonium of this moss dries out, shrinks in diameter, and due to this, the
columella collapses and a high air pressure is resulted inside the capsule. Due to this pressure, the
operculum is thrown away and spores are shot away into the atmosphere. This violent process of
dispersal of spores is known as air-gun mechanism.
Fig. 12.8 A-D Dehiscence of capsule in Sphagnum showing air-gun mechanism of spore dispersal
1. Write an essay on the sporophyte of bryophytes in about 1000 words.

2. Give precise defini ons of sporophyte and spore with reference to bryophytes.
3. “Sporophyte in bryophytes is dependent on gametophyte”. Comment in about 200 words.
4. Sporophyte is the _____ genera on in an alterna on of genera ons.
5. Sporophyte is the genera on producing _____, which are haploid.
6. Sporophyte in bryophytes is usually made up of three parts, viz. _____, seta and capsule.
7. Describe the general structure of sporophyte of bryophytes in about 500 words.
8. Name a bryophyte which lacks foot and seta in its sporophyte.
9. How can you differen ate between seta of liverworts and mosses?
10. The spore-producing organ of the sporophyte of a bryophyte is called _____ .
11. Give an account of development and differen a on of different organs of sporophyte in
bryophytes.
12. What is calyptra?
13. How will you differen ate between haplolepideae and diplolepideae in terms of peristome in
bryophytes?
14. Process of spore dispersal in mosses can be grouped in three broad categories. Name them
and explain them in about 200 words.
13
Spore Germination
and Formation
of Gametophyte
WHAT IS TECHNICALLY A SPORE? 13.1
As explained also earlier, a spore is a “small round cell with a thick wall from which a whole new plant
is produced”. In bryophytes, pteridophytes and spermatophytes, spores are haploid and are produced by
the sporophyte. In bryophytes and pteridophytes, dispersal is achieved by spores. In angiosperms, the
spores develop into small gametophytes in the pollen grains and ovules. In bryophytes and a majority
of other plants, spores are produced as a result of meiosis.
In bryophytes, the spore formation represents the beginning of the gametophytic phase and end
of the sporophytic phase of the life cycle. Because the spore develops due to meiosis in the diploid
spore mother cells of the sporophyte, it is a haploid structure and carries new gene combinations. The
gametophytic plant, produced by spore germination, therefore, has better adaption to the environment
and are genetically uniform.
In brief, it can be mentioned that in bryophytes, the spore is the first cell of the gametophytic
generation. It is a specialised unicellular body which is capable of developing into a new individual.
In the initial stages of their formation, the spores remain in groups of four, thus showing tetrahedral
arrangement. They later on separate into four constituent spores.
ARE BRYOPHYTES HOMOSPOROUS

OR HETEROSPOROUS? 13.2
Plants whose spores are all the same, i.e. possess spores of the same size, are known as homosporous.
On the other hand, plants which produce spores of two different sizes are known as heterosporous.
All bryophytes are usually homosporous, except a few mosses (e.g. Macromitrium salakanum and
Schlotheimia grevillaena) which are heterosporous. In Macromitrium salakanum, the large-sized spores
produce female plants whereas small spores form the male gametophytic plants.
Spore Germina on and Forma on of Gametophyte � 219
SIZE AND OUTPUT OF SPORES 13.3

The size of the spores is highly variable in different bryophytes. Some of the reported range in the
size of the spores of selected genera, as mentioned by Parihar (1987), is from 5m (Dawsonia) to 200m
(Archidium) in Bryopsida, 6 m to 80 m in foliose members of Jungermanniales, and 10 m (Marchantia
polymorpha) to 140 m (Corsinia) in Marchantiales.
The number of spores in each capsule is also highly variable. It is as low as 4 to 28 in Archidium
but as high as 300,000 in Marchantia polymorpha, 400,000 in Diplophyllum albicans, and 1,000,000
in Scapania undulata.
STRUCTURE OF SPORES OF BRYOPHYTES 13.4

A thin wall surrounds the young spore, but soon it becomes thick and bilayered. The inner layer
consists of cellulose and called intine or endosporium, while the outer layer is usually sculpturous
or contains ornamentations and called exine or exosporium. The exine ornamentations may be in the
form of tubercles, spines, folds, ridges, grooves, etc. In many Hepaticopsida, a third spore wall (outer
exosporium or perinium) is also seen. The spores are uninucleate and the nucleus remains surrounded
by granular cytoplasm which contains oil bodies, starch grains, albuminous matter, etc. Chloroplasts are
also present in the spores of several genera including Andreaea, Monoclea, Riccardia, leafy liverworts
and mosses.
SPORE GERMINATION AND FORMATION OF

GAMETOPHYTE IN LIVERWORTS HEPATICOPSIDA 13.5
13.5.1 Marchan ales
Spore germination and formation of the early stages of gametophyte in several members of Marchantiales
has been studied and reviewed by Inoue (1960) and several other workers. On germination, the spore
coat ruptures in many different ways by the enlargement and then emergence of the germ cell. Based
on the polar axis of the spore, Inoue (1960) recognised four types of the spore-coat dehiscence in
Marchantiales. These are (i) irregular, (ii) tangential, (iii) proximal, and (iv) distal. Usually the germ cell
divides by a transverse wall to form a germ rhizoid and a germ tube. But in some genera (e.g. Riccia),
there is no wall formation between the germ cell and germ tube. Seven different patterns of germ
rhizoid formation have been reported in Marchantiales. These are (i) Targionia-type, (ii) Marchantia-
type, (iii) Neohodgsonia-type, (iv) Stephensoniella-type, (v) Reboulia-type, (vi) Mannia-type, and
(vii) Conocephalum-type by Inoue (1960).
The germ tube usually elongates and forms a short filament representing the future gametophyte.
A few chloroplasts and some oil drops are present in the cells of the short filament. Soon, the terminal
cell of the filament divides by two vertical divisions at right angles to one another, to form four cells at
the apex of the filament. This represents the quadrant stage of the young gametophyte. Further vertical
and tangential divisions result in the formation of a several-celled plate of the gametophyte. A distinct
two-sided apical cell is soon differentiated in this multicellular plate. Then there is the differentiation
of the apical cell with four cutting faces. Activity of this apical cell gives rise to a young thallus.
220 � Bryophyta
Inoue (1960) reported five types of the plate formation in Marchantiales. These are (i) Asterella-type,
(ii) Conocephalum-type, (iii) Marchantia-type, (iv) Reboulia-type, and (v) Stephensoniella-type.
13.5.2 Sphaerocarpales
Spore germination of majority of the members of Sphaerocarpales, including Geothallus, Riella and
Sphaerocarpos, has been studied in detail. A slender germ tube emerges and divides by a transverse
division to form a basal cell and a terminal cell. The basal cell does not divide any further and develops
into the first rhizoid. The terminal cell divides by few transverse divisions to form a young, fine filament,
and from this stage onwards the further development of gametophyte is different in different genera.
In Geothallus, the few-celled young filament develops into an erect, unistratose, green, chlorophyll-
containing, flap-like or loose juvenile plant of determinate growth. From the basal part of this juvenile
plant develops the adult plant as a lateral outgrowth. The juvenile plant gives rise to only one adult
plant.
Riella resembles Geothallus in producing a single erect, unistratose juvenile plant of determinate
growth but it is comparatively larger than that of Geothallus. At the base of this juvenile plant, a series
of cells remains embryonic and function as its intercalary meristem. This meristem first makes the
juvenile plant more active by adding new cells. Lower and upper regions are soon differentiated in
this young gametophyte due to the activity of this intercalary meristem. Soon the young gametophyte
attains a size of 5 to 8 cm or more.
In Sphaerocarpos, the slender germ tube emerges, pushes through a slit on the outer face of the
spore, divides transversely into a terminal cell and a basal cell, and a filament develops due to some
transverse devisions in the terminal cell. Cells of this filament divide and form a germinal disc at
right angles to a short multiseriate filamentous body. This multicellular germinal fisc transforms into a
juvenile thallus. A lateral outgrowth develops from this juvenile thallus and changes into an adult plant
of Sphaerocarpos.
13.5.3 Jungermanniales
The young plant, formed after the early divisions of spore in Jungermanniales, has been termed sporeling.
The so-called sporeling (Fulford, 1956) includes all the stages of a young developing plant, such as
(i) protonema, (ii) the shoot with primary leaves, and (iii) shoot with underleaves and juvenile stages.
The protonema stage, as explained by Fulford (1956) includes “all stages from the first division
of the spore up to the formation of an apical cell with three cutting faces by which the leafy shoot
is formed”. He divided all Jungermanniales into the undermentioned two groups, which include ten
different types of sporelings. Two types are included under Group A and the remaining eight types
are included under Group B. In Group A, protonema develops outside the exospore, while in Group
B, the protonema develops completely or at least partially, within the stretched exospore. In Group A,
the juvenile leaves are with a deep sinus and spreading lobes, but in Group B, the juvenile leaves are
large and saccate inflated.
Group A
(a) Cephalozia Type It contains a simple or branched filamentous protonema with stems at the tips of
branches, e.g. Cephalozia bicuspidata (Fig. 13.1.A).
(b) Nardia Type It contains a globose, multicellualr protonemata, e.g. Nardia scalaris (Fig. 13.1B).
Group B
(a) Radula Type It contains a disciform protonema, e.g. Radula complanata (Fig. 13.1 C).
(b) Frullania Type It contains a multicellular (up to 50 or more cells), globose protonema within the
exospore, e.g. Frullania dilatata (Fig. 13.1 D).
(c) Lopholejeunea Type It contains a globose protonema made up of only a few cells, e.g. Archilejeunea
(Fig. 13.1 E).
(d) Leucolejeunea Type It contains elongate cylindrical protonema, as in Leucolejeunea (Fig.

13.1F).
(e) Lejeunea Type It contains a narrow, unistratose protonema which develops due to the activity of
an apical cell with two cutting faces, as in Lejeunea (Fig. 13.1 G).
(f) S ctolejeunea Type It contains unistratose protonema which does not grow by an apical cell, as in
Stictolejeunea (Fig. 13.1 H).
(g) Ceratolejeunea Type It contains a bifold protonema made up of two thalloid stages, as in
Ceratolejeunea (Fig. 13.1 I).
(h) Unnamed type of bifold protonema with a multicellular primary protonema and a ribbonlike
secondary protonema.
Fig. 13.1 A-I Various types of protonema of bryophytes. A, Cephalozia; B, Nardia; C, Radula; D, Frullania;
E, Lopholejeunea, F, Leucolejeunea; G, Lejeunea; H, S ctolejeunea, I, Ceratolejeunea
222 � Bryophyta
SPORE GERMINATION AND FORMATION OF GAMETOPHYTE

IN HORNWORTS ANTHOCEROPSIDA 13.6
Spore germination in Anthoceros (Fig. 8.9A-H) starts by rupturing of the exospore along the triradiate
ridge, the endospore protrudes as a papilla and forms a germ tube. The plastid, oil globules and other
food materials of the spore pass into the germ tube. Initial two divisions in the germ tube are transverse,
forming a 3-celled short filament (Fig. 8.9 D, E). A cylindrical elongated young gametophyte is resulted
soon. For more details, refer to Article 8.5.14 (Fig. 8.9 A-H).
In Notothylas, a spore germinates to form a mass of cells which is known as a germ disc. It is made
up four or eight cells. One or two basal cells of the germ disc extend to form rhizoids. They are not
separated by any wall. A marginal meristem soon differentiates and initiates the growth of the thallus.
SPORE GERMINATION AND FORMATION OF

GAMETOPHYTE IN MOSSES BRYOPSIDA 13.7
Bryopsida usually possess a well-branched filamentous protonema, from which develop numerous buds
(Fig. 13.2 A-G). Thalloid protonema is rarely seen amongst Bryopsida, except Andreaea, Sphagnum,
Tetraphis and a few more members.
Each bud produced on the filamentous protonema shows differentiation of a tetrahedral apical cell.
It divides and redivides to form a leafy shoot system of gametophore. Protonema of most mosses has
a heterotrichous structure, of which the prostrate multicellular filaments creep on the substratum, and
from these filaments develop lateral erect filaments.
13.7.1 Forma on of Buds

A bud develops as a lateral protrusion from a cell of the parent filament. Usually, it develops just behind
a septum. It first divides transversely to form one or two stalk cells and an upper cell. The upper cell
swells, divides by three successive oblique divisions to form a future gametophore. Usually, a bud
develops near the base of the parent filament of the aerial erect system, but variations occur commonly
in different mosses. Both prostrate as well as erect green branches bear buds in many mosses including
Ceratodon, Mnium and Pohlia.
13.7.2 Requirements for Bud Forma on in Mosses

In the normal conditions, certain minimum light intensity and sugar requirements are essential for
bud formation. It has been experimentally proved that kinetin “acts as an agent which creates centres
of attraction in moss protonema” (Bopp, 1963). Adequate nutrients and certain growth substances are
also the major requirements for bud formation in Pohlia nutans, Tortella caespitosa and some more
mosses. It has been proved experimentally by various workers that there is a definite role of light
for the germination of spores in bryophytes (Valanne, 1971; Cove et al. 1978, Schield, 1981). Low
temperatures are helpful in germination of liverwort spores. For more details of the role of temperature
in spore germination, consult Steiner (1964), and Chopra and Sood (1973).
D
B C
A
Protonema
Bud
Leaves
Rhizoids
F
Stem
Bud
Secondary protonema
Rhizoid G
Gemma
Fig. 13.2A-G Spore germina on in Funaria hygrometrica
13.7.3 Protonema of Some More Mosses

In Buxbaumia aphylla (Fig. 13.3A), the protonema is persistent, green, filamentous and much branched.
The leafy gametophores arising from it are small. Persistent protonema also develop in some more
genera including Pogonatum aloides, Nanobryum dummari and Schistostega pinnata.
In Ephemeropsis tjibodensis (Fig. 13.3B), the protonema is quite extensive while the gametophore
is very small. From the flanks of the main axis of this moss shoot out anchoring organs called hapteres.
Well-branched forked green assimilatory short shoots develop from the dorsal surface.
224 � Bryophyta
Fig. 13.3 Protonema of Buxbaumia aphylla (A) and Ephemeropsis (B)
SPORE VIABILITY 13.8

Spores of liverworts remain viable for a few weeks to a few months while that of mosses remain
viable for several years. Amongst liverworts also, the foliose or leafy liverworts growing in mesic
environment have a shorter period of viability while the thalloid liverworts, growing comparatively
in xeric environment have a longer period of viability for many months. Looking over the mosses,
the spores of Sphagnum remain viable from two to three years while that of Funaria hygrometrica
remain viable for 10 to 12 years, and that of Oedipodium remain viable for as much as 20 years or even
more.
POLARITY IN SPORE GERMINATION 13.9

Polarity means existence of a definite axis. Bryologists (Bentrup, 1968); Schulz, 1972; Olesen and
Mogensen, 1978; Brown and Lemmon, 1980) have worked on polarity in bryophytes. It has been
conclusively established that spore germination in bryophytes show polarity. Spores of Marchantiales
and some other investigated thalloid bryophytes are polar structures. On the other hand, spores of
mosses are apolar structures, and they generally do not show polarity in their spores. According to
Brown and Lemmon (1980), however, the subcellular structure of moss spore shows an innate polarity.
The term “innate” actually refers to behaviour which normally occurs in all members of a species
despite natural variation in environmental influences.
1. Give a precise defini on of spore.

2. In bryophytes, pteridophytes and _____, the spores are haploid and are produced in the
_____.
3. In bryophytes, the spores are produced as a result of which division: mitosis or meiosis?
4. The spore forma on in bryophytes represents the beginning of _____ phase.
5. In bryophytes, the spores are always which type of structures: diploid or haploid?
6. What is the first cell of a gametophy c genera on in bryophytes?
7. Write an essay on spore germina on and forma on of gametophyte in bryophytes in about
500 words.
8. Leaving aside only a few excep ons, are all bryophytes homosporous or heterosporous?
9. Write a brief scien fic note on size and output of spores.
10. Size of the spore of which of the following bryophytes is smallest?
(a) Dawsonia, (b) Marchan a polymorpha, (C) Corsinia, (d) Archidium
11. Give an account of spore germina on and forma on of gametophyte in liverworts.
12. What is protonema? Describe it with par cular reference to Jungermanniales.
13. Describe spore germina on and gametophyte forma on in Bryopsida (mosses).
14. Write scien fic notes in about 100 words each on the following:
(a) Spore viability in bryophytes
(b) Polarity in spore germina on in bryophytes
14
Factors Affecting
Sexuality in
Bryophytes
MAJOR FACTORS AFFECTING SEXUALITY 14.1
Besides widespread occurrence of vegetative reproduction, the bryophytes also reproduce sexually,
provided necessary climatic conditions are available in their surroundings. Sexual reproduction in any
group of plants or animals is also necessary and essential because this is the only way for genetic
variability, which is required for existence and evolution of the group.
In bryophytes, initiation of gametangia, their development and appearance of sporophytes are
influenced by a variety of factors including (i) photoperiod, (ii) temperature, (iii) carbohydrates, (iv)
nitrogenous substances, (iv) pH and hydration of medium, and (iv) growth regulators. All these factors
affect sexuality in bryophytes, and are discussed briefly here in this chapter.
INFLUENCE OF PHOTOPERIOD ON SEXUALITY 14.2

“The number of hours of daylight needed by a plant before it will begin to flower” is called photoperiod,
and this phenomenon of the “physiological responses of an organism to changes in the lengths of day
and night is called photoperiodism”. The photoperiodic control of sexuality in bryophytes was first
studied by Wann (1925), and these details were published in the American Journal of Botany. In
several further studies of different workers, it was established that photoperiodic control of gametangial
initiation is more prevalent amongst Hepaticopsida and Anthoceropsida than Bryopsida or mosses. As
we see commonly among angiosperms, in bryophytes also there exist long-day plants, short-day plants
and also day-neutral plants. Differing from angiosperms, however, the photoperiod in bryophytes refers
to “hours of duration of light during 24-hour cycle” and it does not refer to length of the dark period,
which is actually very critical for control of flowering in higher plants.
14.2.1 Long-day Bryophytes

Wann (1925) was the first to report Marchantia polymorpha as a long-day bryophyte. During long
days, it grows more vigorously and becomes fertile by producing gametangia. During short days, only
Factors Affec ng Sexuality in Bryophytes � 227
a few or no gametangia are produced by this liverwort. Some other bryophytic taxa which have been
reported by different workers as long-day plants are Conocephalum conicum, Diplophyllum albicans,
Lophocolea cuspidata, Pellia epiphylla and Riccardia multifida. All these produce gametangia only
when they receive 16–18 hours of light per day.
Proskauer (1967) confirmed experimentally that a hornwort (Phaeoceros laevis) from western
Himalayas is a long-day bryophyte. It grows luxuriantly and produces gametangia only during long
days.
Amongst the long-day bryophytes, temperature also plays a definite role besides light intensity for
initiation of gametangia. Pellia epiphylla, a long-day bryophyte, produces more gametangia at higher
temperatures. Marchantia polymorpha becomes more fertile during long days when the temperature
is above 20°C, and it produces only a few or no gametangia during long days when the temperature is
below 10°C. Dua et al. (1982) proved Riccia gangetica, a common liverwort of Indo-Gangetic plains,
to be a long-day bryophyte.
Plagiomnium undulatum, a moss of Britain, is a long-day plant and produces male and female
gametangia only during high intensity of daylight of 15 to 18 hours per day. Some other mosses which
fall under the category of long-day plants are Barbula gregaria, Bryum argenteum and Bartramidula
bartramioides.
14.2.2 Short-day Bryophytes

Asterella tenella and Targionia hypophylla of Hepaticopsida; Anthoceros laevis, Notothylas
orbicularis and Phaeoceros laevis of Anthoceropsida; and Sphagnum subnitens of Bryopsida are some
examples of short-day bryophytes. According to Bapna et al. (1984), Targionia hypophylla produced
archegoniophores only under cool short-day conditions. Mosses are seldom short-day plants, except a
few, e.g. Sphagnum subnitens.
14.2.3 Day-Neutral Bryophytes

A majority of mosses and only a few liverworts are day-neutral bryophyte, and in all such genera,
the gametangial initiation is independent of the length of the day. Amongst the mosses, common
day-neutral members are Funaria hygrometrica, Laptobryum pyreforme, Physcomitrium pyriforme
and Polytrichum aloides. Cryptothallus mirabilis of Metzgeriales (Hepaticopsida) is a day-neutral
bryophyte, in which initiation of gametangia is independent of the length of the day. In this member of
Metzgeriales, the initial requirements of the initiation of gametangia are cold environment and then rise
in temperature up to 21°C and there is not much role of the day length.
INFLUENCE OF TEMPERATURE ON SEXUALITY 14.3

As mentioned earlier also under Article 14.2, besides photoperiod, temperature also influences and
controls the initiation and development of sexuality in bryophytes. There are, however, many bryophytes
which are not sensitive to temperature and form sex organs in a wide range of temperature from very
low to very high.
228 � Bryophyta
On the basis of the influence of temperature on sexuality, all bryophytes may be grouped under the
following five categories:
1. A cold-conditioning of some of the bryophytes is a prior condition for the gametangial
initiation.
2. Formation of gametangia in some bryophytes is possible only at low temperature. Higher
temperatue in such bryophytes inhibits gametangial formation.
3. Formation of gametangia in some bryophytes is possible ony at fairly high temperature.
4. Formation of gametangia in some bryophytes is inhibited at low temperature and possible only
at temperatures above 21°C.
5. Formation of gametangia takes place over a wide range of temperatures, i.e. these bryophytes
are insensitive to temperature.
In Cryptothallus mirabilis and Lunularia cruciata, gametangia are formed at 20–21°C but both these
liverworts have an obligate requirement of cold-conditioning of their thalli at 4–10°C as a stimulus. In
Funaria hygrometrica, gametangia are formed when day and night temperatures are maintained at 10°C
and 7°C, respectively. Plants of this common moss remain sterile when day and night temperatures are
at 5°C and 3°C and also 17°C and 15°C, respectively. According to Benson-Evans (1964), Riccia
glauca in Britain is a short-day bryophyte and forms gametangia when day and night temperatures
are 18°C and 10°C, respectively. Its plants remain sterile at 21°C. However, Chopra and Sood (1973)
reported that gametangia formation in Riccia crystallina in India is at its peak at a temperature range
of 8 to 15°C. Kumra and Chopra (1984) also demonstrated the temperature requirements for initiation
of gametangia as 20°C in Philonotis turneriana. In Leptobryum pyreforme, the plants remain sterile at
25°C, but at 20°C only a few plants become fertile. Early gametangial initiation in very large number
of plants in L. pyreforme is observed when the temperature is as low as approximately 10°C. It shows
protrandrous condition also, i.e. antheridia develop first and archegonia later on. Benson-Evans
(1964) demonstrated that the moss Polytrichum aloides and two liverworts (Marchantia polymorpha
and Conocephalum conicum) show their specific temperature requirements of gametangial initiation
as 21°C. All these three bryophytes remain sterile at low temperatures. Some mosses (e.g. Barbula
gregaria, Bartramidula bartramoides and Bryum argentatum) and liverworts (e.g. Pellia epiphylla)
develop gametangia under a wide range of temperature.
INFLUENCE OF CARBOHYDRATES ON SEXUALITY 14.4

Wann (1925), while studying sexuality in bryophytes, observed that in Marchantia polymorpha the sex-
organ initiation takes place at a higher carbohydrate/nitrogen ratio. Rao and Das (1968) also confirmed
this finding in some other bryophytes including Plagiochasma articulatum, Reboulia hemispherica and
Fimbriaria angusta.
Excess of sugar shows inhibitory effect on sexuality in several Hepaticopsida members including
Athalamia pusilla, Fossombronia himalayensis, Riccia crystallina and R. frostii. Even in the absence
of carbohydrate supply, the gametangial formation takes place in some mosses, e.g. Barbula gregaria
and Bryum argenteum. However, Dua et al. (1982) opined that sucrose is more effective for initiation
of sexuality in Riccia gangetica and R. crystallina.
INFLUENCE OF NITROGENOUS SUBSTANCES ON SEXUALITY 14.5

It has been proved experimentally that in liverworts, inorganic nitrogen proves inhibitory for
gametangial formation while organic nitrogen favours it. Sood (1974) proved experimentally in Riccia
crystallina that if organic nitrogen is supplied in the form of either urea, casein hydrolysate or yeast
extract, it favours formation of archegonia. Almost similar results were seen in Riccia frostii and
Pohlia nutans by other workers. Antheridial formation, however, is not affected by the supply of these
nitrogenous substances. Woodfin (1976) proved that in several species of Riccia, including R. flutians,
the gametangial formation stops if concentration of ammonium nitrate is reduced to half.
INFLUENCE OF pH AND HYDRATION OF

MEDIUM ON SEXUALITY 14.6
pH is actually the negative of the logarithms of hydrogen-ion concentration in aqueous solution. Low
pH is acid while high pH is alkaline. pH of 7 is neutral. pH and hydration of medium show definite
effect on sexuality in bryophytes. A few such examples are listed below:
1. In Sphaerocarpos donnellii, male plants grow well in alkaline medium or near-neutral medium
whereas female plants grow well in low pH.
2. Dua (1983) observed that in Riccia gangetica, the maximum antheridial formation is seen at a pH
of 4.5 while initiation of archegonia is optimum in alkaline pH of 7.5
3. Rahbar and Chopra (1982) observed that in Bartramidula bartramioides, a moss, the optimum
negative growth takes place at a pH of 4.5, and as the pH increases, the vegetative growth declines. The
optimum initiation and development of gametangia in this moss is seen at a pH of 7.5.
4. Sood (1974) observed in Riccia crystallina that level of mineral elements and hydration of
medium affect the initiation of gametangia in this common Hepaticopsid. If level of mineral elements
and hydration of medium is reduced, it favours maleness. On th the other hand, higher levels favour
femaleness.
5. Vashistha (1985) observed that female thalli of Riccia frostii remain sterile in liquid medium
while they start producing archegonia in agar cultures.
6. Differing from this trend of R. frostii, Kumar (1981) observed that in mosses (e.g. Barbula
gregaria), the initiation of gametangia is delayed in the presence of agar, and an increased response of
gametangial initiation is observed in liquid medium.
INFLUENCE OF GROWTH REGULATORS ON SEXUALITY 14.7

Almost all categories of hormones (auxins, gibberellins, cytokinin) have been reported to promote
initiation of sexuality in bryophytes. They are effective individually as well as in combination.
230 � Bryophyta
14.7.1 Auxin
“Auxin” is actually a general name for an important group of plant hormones. Indole Acetic Acid
(IAA) is the most common auxin. Auxins have been reported to favour femaleness in liverworts. If IAA
is applied on the vegetative thalli of Marchantia polymorpha, it results into the formation of structures
similar to the receptacles. Almost similar results are seen if the auxin 2, 4-D (2, 4-Dichlorophenoxyacetic
acid) is applied. (2, 4-D is a “synthetic auxin used as a selective herbicide and in media for tissue
culture”). Rao and Das (1968) observed initiation of archegonia by increasing the level of IAA in
several bryophytes including Asterella angusta, Pallavicinia canarus, Plagiochasma articulatum and
Reboulia hemispherica. Chopra and Sood (1973), Kumra and Chopra (1984), and Vashistha (1985)
observed that auxins favour femaleness in several species of Riccia, including R. crystallina, R. frostii
and R. gangetica. Much work has not been done on the role of auxins in the initiation of sexuality in
bryophytes. Chopra and Kumra (1983) observed that IAA stimulates the formation of antheridia in two
mosses, viz. Barbula gregaria and Bryum argenteum.
14.7.2 Gibberellins
Gibberellins are a group of chemically complex plant hormones, important in control of tropisms,
the lengthening of cells during growth, germination and other processes. Gibberellins promote
maleness in liverworts as well as in mosses. Chopra and Kumra (1983), and Sarla (1986) reported
antheridial induction at very low level of gibberellic acid in Philonotis turneriana and Riccia discolor,
respectively. In basal medium, both these bryophytes remain sterile. Chopra and Sood (1973) reported
that in monoecious Riccia crystallina, gibberellic acid enhanced antheridial formation remarkably.
Chopra and Kumra (1986) reported that the number of antheridia increased at all levels of GA3 in
R. gangetica. In yet another study, Chopra and Gupta (1992) observed that gibberellin suppresses
formation of archegonia in female clones of Riccia discolor.
14.7.3 Cytokinins
Cytokinins are a group of plant hormones (e.g. kinetin), which control cell division. They support
femaleness in some hepaticopsids and mosses. Chopra and Sood (1973) observed that in monoecious
Riccia crystallina, kinetin increased the archegonia production without showing any effect on the number
of antheridia. Kumra and Chopra (1984) also observed almost the same results in Riccia gangetica.
Vashistha (1987) reported in Riccia frostii that application of cytokinins also promotes formation
of archegonia in female plants. Chopra and Rawat (1977) studied that cytokinins show no effect on
formation of sex organs in monoecious species of the moss Leptobryum pyriforme. Chopra and Kumra
(1983) reported that cytokinins are ineffective on male clones of mosses, such as Bryum coronatum
and Barbula gregaria. As reported by Chopra and Mehta (1987), cytokinin increased the frequency of
fertile gametophytes in male clones of Microdus brasiliensis. In mosses, such as Bryum argenteum, a
mixture of auxin and cytokinin promotes formation of both male and female gametangia.
1. Make a list of at least five factors which affect sexuality in bryophytes.

2. Describe influence of different factors affec ng sexuality in bryophytes.
3. Define the term ‘photoperiod’ and describe its role on sexuality in bryophytes.
4. Name at least three long-day bryophytes.
5. State whether a majority of mosses are day-neutral bryophytes or long-day bryophytes?
6. “Mosses are seldom short-day plants”. Is it true or false?
7. Write a detailed note on the influence of temperature on sexuality in bryophytes.
8. Is pH of 7 acidic, alkaline or neutral?
9. Describe in detail the role of growth regulators on sexuality in bryophytes in about 200
words.
15
Alternation
of Generations
WHAT IS ALTERNATION OF GENERATIONS? 15.1
“The life cycle of bryophytes, pteridophytes and spermatophytes, which consists of a haploid
gametophyte producing gametes followed by a diploid sporophyte producing spores” is known as
alternation of generations. It is actually the regular alternation of two types of individuals in the life
history of plants or animals. Typically, in plants, there is alternation of a diploid sporophyte and a haploid
gametophyte. If the two generations (i.e. sporophyte and gametophyte) are morphologically similar,
it is known as isomorphic alternation of generations, but if two generations are morphologically
different, then it is known as heteromorphic alternation of generations.
In bryophytes, the dominant generation is the gametophyte (Fig. 15.1), and it is markedly different
from the sporophyte, and hence these plants show heteromorphic alternation of generations. The
gametophyte or gamete-producing generation has a single set of genomes while the sporophyte or
Fig. 15.1 Diagramma c representa on of alterna on of genera ons

Alterna on of Genera ons � 233
spore-producing generation is morphologically different from the gametophyte and is diploid. The
sporophyte undergoes meiosis to form haploid spores which germinate to produce haploid gametophytes
(n). The gametophyte undergoes gametogenesis and produces haploid male and female gametes
(antherozoids and egg) that fuse sexually to form a diploid zygote, from which develops the diploid
sporophyte.
WHO DISCOVERED ALTERNATION OF GENERATIONS

IN ARCHEGONIATAE? 15.2
“Archegoniatae” includes the plants having archegonia, applied to bryophytes, pteridophytes and
gymnosperms. Technically speaking, an “archegonium” is the female sex organ of bryophytes,
pteridophytes and gymnosperms, containing the egg inside a cellular jacket. W. Hofmeister (1851) was
the first to show that life cycle of members of Archegoniatae includes two distinct and well-marked
generations: (i) the gametophyte, which contains gamete-producing sex organs, i.e. archegonia and
antheridia, and (ii) the sporophyte, which produces the asexual spores; and both these generations (i.e.
gametophyte and sporophyte) appear one after another in regular alternation succession to complete
the life-cycle, and this entire cycle represents alternation of generations. Hofmeister’s 1851 findings
were published in the book Vergleichende Untersuchungen.
WHAT ARE ANTITHETIC ALTERNATION AND HOMOLOGOUS

ALTERNATION OF GENERATIONS? 15.3
It was L. Celakovsky (1874) who coined these two terms (antithetic and homologous) in connection
with alternation of generations, and explained them as under
1. Antithetic means heteromorphic. The term describes the life cycle in which the alternating
generations are morphologically and physiologically distinct, as in bryophytes and
pteridophytes. Celakovsky (1874) explained it as “antithetic alternation of two generations
phylogenetically distinct, i.e. where a new stage (sporophyte) has been interpolated between
pre-existing generations (gametophytes)”. According to him, this type of alternation might
have probably developed” independently in several different groups, but was most evident in
Archegoniatae” (Celakovsky, 1874).
2. Homologous means isomorphic. The term describes the life cycle in which alternating
generations are morphologically identical, as commonly occurs in algae (e.g. Dictyota) and
fungi. Individuals may be identified as sporophyte or gametophyte by observation of the
reproductive process. Such life cycles are seldom seen in archegoniate plants. Celakovsky
(1874) explained it as “homologous alternation of two or more generations phylogenetically
similar to one another, but differing in the presence or absence of sexual organs.”
PHENOMENA OF APOGAMY AND APOSPORY 15.4

In connection with nature of alternation of generations and origin of sporophytes, two phenomena were
later on recognised. These were termed apogamy and apospory.
234 � Bryophyta
1. Apogamy Apogamy was first discovered by Farlow in 1874 but the name (apogamy) to this
phenomenon was given by De Bary in 1874. Apogamy is a phenomenon of asexual reproduction in
which embryos and propagules are produced without the occurrence of meiosis. Thus, the sporophyte
might develop vegetatively from the gametophytic tissue without the process of fertilization.
2. Apospory Apospory was first discovered by Pringsheim in 1876. Apospory is a phenomenon

of production of a diploid gametophyte from the vegetative cells of the sporophyte, i.e. without the
production of spores.
ANTITHETIC THEORY 15.5

It is also called interpolation theory or intercalation theory. As mentioned earlier also, the antithetic
theory was first proposed by Celakovsky in 1874. According to this theory, the “gametophyte (or sexual)
generation is original or historically a prior generation, while the sporophyte (or spore-producing)
generation” is completely a new phase. This new sporophytic phase is “derived from the progressive
elaboration of the zygote of some algal ancestor”. This phase is also intercalated or interpolated into the
life cycle, during the course of evolution, “between the successive events of fertilization and meiosis,
and is therefore different in structure from the gametophyte from its very inception” or right from the
very beginning.
FO Bower (1935) has been the main supporter of antithetic theory of Celakovsky (1874) besides
many others, including E Strasburger (1894), F Cavers (1911) and DH Campbell (1940). According
to this theory of alternation of generations, the sporophyte originated from the zygote (or unicellular
sporophyte) of some ancestor of filamentous green algae, e.g. Oedogonium. From this simple unicellular
sporophyte or zygote, the complex sporophyte evolved gradually by progressive elaboration, of course,
through several undermentioned stages:
1. First Stage The life cycle in the earliest hypothetical land plants would be resembling to that
of many filamentous green algae, e.g. Oedogonium (Fig. 15.2A), in which the sporophyte would be
represented by unicellular diploid zygote. It divided meiotically to form four haploid zoospores, also
called zoomeiospores (Fig. 15.2B-E).
2. Second Stage Coleochaete-like (Fig. 15.3) green algal members represented this stage, in
which the zygote increases greatly in size and divides meiotically to form 16–32 biflagellate swarmers
or zoospores. Reduction division in this genus takes place in the first dividion of the diploid zygote, and
thus the contrasting type of Coleochaete individual, formed from the zygote, is haploid in nature.
3. Third Stage (Simplest Sporophyte) In this next stage, the zygote is retained within the
archegonium and did not divide meiotically. It first divided mitotically to form a diploid multicellular
body of spore-producing cells. Each such spore-producing cell is diploid, divided meiotically to form
four haploid spores. Although hypothetical, but this would have been the simplest type of multi-
cellular sporophyte of plants possessing multicellular embryo (i.e. Embryophyta, which includes
bryophytes, pteridophytes and spermatophytes). In such a sporophyte, all the cells were sporogenous.
However, it differs in structure from the gametophyte.
Fig. 15.2 A-E Oedogonium. A, A filament of monoecious species; B-C, Zygotes present inside and outside
the oogonium; D, Showing four daughter protoplasts formed a er meiosis; E, Showing haploid
zoomeiospores
236 � Bryophyta
Fig. 15.3 Coleochaete divergens
4. Fourth Stage This stage is seen in Riccia (Fig. 15.4 A) in which the diploid zygote divided
and redivided to form a multicellular, spherical diploid body, of which the outermost layer became
sterile and formed the jacket while the inner central mass remained sporogenous and diploid in nature.
Now, reduction division took place in each cell of the diploid sporogenous tissue to form four haploid
spores. There was no differentiation of the foot, seta and capsule in this simple stage of sporophyte.
Its growth was also limited, and it remained surrounded by a single-layered jacket. A very large part is
thus capable to form the spores.
According to F O Bower, it was this above-mentioned type of simple sporophyte of Riccia, from
which evolved the more complex types of sporophytes of Embryophyta (i.e. bryophytes, pteridophytes
and spermatophytes), and all this happened by “progressive sterilization of potentially sporogenous
tissue”. A series can be traced amongst different members of bryophytes which shows (i) ever-
increasing sterilization of sporogenous tissue, and (ii) such a sterilized tissue shows its regular diversion
to somatic functions. From simple sporophyte of Riccia developed most complex type of sporophytes
of mosses, e.g. Funaria and Polytrichum, via a series of different other members of bryophytes.
5. Fi h Stage In Riccia crystallina and Oxymitra, a stage slightly ahead of Riccia is seen of “pro-
gressive sterilization of potentially sporogenous tissue.” Both these contain a simple sporophyte made
up of a single-layered sterile jacket which encloses a mass of sporogenous tissue. But some of the
potential spore mother cells remain unable to produce spores. They instead form absorptive nutritive
cells. These cells have been considered forerunners of true elaters.
6. Sixth Stage It is seen in genera such as Sphaerocarpos (Fig. 15.4 B) and Corsinia. The entire
basal part of the sporophyte gets sterilised in these members in the form of a small sterile foot made of
only a few cells in Corsinia, or a small bulbous foot and a 2-cells broad, narrow seta in Sphaerocarpos.
A single-layered sterile jacket also surrounds the capsule in both Corsinia and Sphaerocarpos, as in
the fourth and fifth stages described above. Sterile nurse cells are also present in the Corsinia capsule.
These nurse cells are homologous with elaters but lack their characteristic thickenings. Presence of
capsule at the apex and a foot at the base shows the existence of polarity amongst both Corsinia and
Sphaerocarpos.
Fig. 15.4 Evolu onary series of sporophytes in bryophytes. A, Riccia; B, Sphaerocarpos s pitatus;
C, Targionia hypophylla; D, Marchan a polymorpha; E, Pellia epiphylla
238 � Bryophyta
Pseudoelaters
Spores
Jacket
Columella
Jacket
Spore tetrad
Columella
Archesporium
Meristematic
region
Foot
Thallus cells
Fig. 15.5 Longitudinal sec ons of different stages of sporophyte of Anthoceros
7. Seventh Stage Targionia (Fig. 15.4C) shows this next stage of evolutionary series of sporo-
phytes, in which more sterile region is present than in the sixth stage. The sterile region includes a
broader foot, a more developed and long seta, a single sterile jacket layer of jacket of the capsule and
about half of the sporogenous cells becoming sterile in the form of several elaters, each with 2 or 3
spiral thickenings.
8. Eighth Stage Marchantia polymorpha (Fig. 15.4D) shows the next stage of evolution among
sporophytes of bryophytes, in which more sterile tissue is present in the form of (i) broad foot, (ii) more
developed, long and thick seta, (iii) single-layered sterile jacket of capsule, (iv) sterile cells in the form
of cap at the apex of the capsule, and (iv) long and well-developed large number of spirally thickened
elaters.
9. Ninth Stage In Pellia epiphylla (Fig. 15.4E) and Riccardia, sterilization of potentially sporog-
enous tissue is more developed than the above-mentioned eighth stage of Marchantia. In both these
genera, the sterile tissue consists of a large foot, more developed seta, two to many-layered sterile
jacket around the capsule, diffused elaters and a sterile mass of cells in the form of elaterophore, either
at the basal end of the capsule (Pellia, Fig. 15.3E) or at the apical end of the capsule (Riccardia). Only a
very small percentage of actual sporogenous tissue is present in the sporophyte in the form of spores.
10. Tenth Stage More sterilization of potential sporogenous tissue is seen in this stage, exempli-
fied by Anthoceros (Fig. 15.5). The sterile tissue in the sporophyte includes (i) a massive foot, (ii) a 4
to 6-layered capsule wall, (iii) a central column of elongated cells in the form of columella, and (iv)
pseudoelaters. The sporophyte in Anthocerotoes becomes more independent due to the presence of
chlorophyllous tissue, an epidermal layer and presence of stomata in the epidermis.
11. Eleventh Stage In Funaria hygrometrica (Fig. 15.6), Polytrichum and some other higher
bryopsida, there is seen the height of the sterilization of the potentially sporogenous tissue. The sterile
tissue is present in the form of a large foot, very long seta, apophysis, multilayered capsule wall, per-
Operculum Peristome
Jacket
Columella
Air space
Archesporium
Apophysis
Stomata
Seta
Fig. 15.6 Longitudinal sec on of the capsule of Funaria hygrometrica

240 � Bryophyta
istome, operculum, a well-developed columella and also aerenchyma and wall around the spore sac.
The potential sporogenous tissue is present only in the form of spores in the spore sac.
During course of time there was seen more elaboration of the sterile vegetative structure of the
sporophyte, and due to the development of roots and leaves, the sporophyte became independent and
achieved a dominant phase of the life cycle. Bower put forward the view that origin of alternation of
generations is also related with a change in habit from aquatic to subaerial life or terrestrial life.
FINAL PICTURE ABOUT NATURE AND EVOLUTION

OF PRIMITIVE SPOROPHYTE 15.6
15.6.1 View of Bower and his Supporters
Botanists such as Bower (1908), Cavers (1910), Campbell (1940) and several of their supporters
believe that the most primitive sporophyte is the simplest sporophyte of Riccia, and from this simplest
sporophyte evolved the more advanced type of sporophytes by progressive sterilization of potentially
sporogenous tissue. This all happened through various stages discussed above under “Antithetic
Theory” (Article 15.5).
15.6.2 View of Kashyap and his Supporters

Shiv Ram Kashyap (1919) and his supporters (Church, 1919; Evans, 1939 and others) believe
differently, and opined that “the simplest sporophyte of Riccia is not a primitive type, but it is a reduced
type evolved as a result of descending or regressive evolution”, to which they called “progressive
simplification”. It was AH Church (1919), who proposed his theory that “the hypothetical ancestral
sporophyte of Bryophyta was an erect leafy shoot, which was independent.” During the process of
descending or regressive evolution, this erect leafy sporophyte passed through the following changes:
1. It attached permanently to the gametophyte.
2. Its leaves lost because of desiccation and intense isolation.
3. It became more dependent on the gametophyte because of the disappearance of air spaces in
its photosynthetic system.
4. Its chlorophyll-containing tissue became reduced.
5. Its stomata became functionless (e.g. Sphagnum).
6. After becoming functionless, the stomata disappeared completely (e.g. Marchantiales).
Church (1919) thus opined that sporophytes of Bryopsida (e.g. Funaria, Polytrichum) and
Anthocerotopsida (e.g. Anthoceros) are (i) “primitive and nearer to the ancestral type”, (ii) “markedly
reduced and simplified”, as that of Marchantiales (e.g. Marchantia), and (iii) “highly reduced”, as in
Riccia.
HOMOLOGOUS THEORY 15.7

15.7.1 What is Homologous Theory?
It is also called “modification theory” or “transformation theory”, and it was N Pringsheim (1876,
1878) who proposed this theory. According to this theory, “the sporophyte and gametophyte generations
are fundamentally similar in nature and the sporophyte is a direct modification of the gametophyte
and is not a new structural type” (Pringsheim, 1876, 1878). The homologous theory of Pringsheim
got support from several eminent botanists, including Zimmermann (1930), Bold (1938) and Fritsch
(1945).
15.7.2 Some Evidences to Support Homologous Theory

Listed under are the evidences which support the homologous theory of alternation of generations:
1. Existence of isomorphic alternation of generations in a large number of algae belonging to
Cladophorales, Chaetophorales, Ulvales, Dictyotales and Ectocarpales.
2. Existence of photosynthetic tissue in sporophytes of several members of Bryopsida (e.g.
Funaria), Anthoceropsida (e.g. Anthoceros), etc., which shows the nature of self-sufficiency
of both gametophyte and sporophyte of these members.
3. Similarity in structure between the primitive gametophytes (e.g. Psilotum, Tmesipteris) and
primitive sporophytes (e.g. Rhynia, Psilotum) of many members of pteridophyta.
4. Similarity in a large number of bryophytes and pteridophytes in the occurrence and type of
phenomena like apogamy and apospory.
GENETICAL AND ONTOGENETIC VIEWS ON

ALTERNATION OF GENERATIONS 15.8
Stebbins (1950), while working on variation and evolution of plants, studied some genetic aspects of
alternation of generations, and proposed that “homologous theory seems more agreeable than antithetic
theory”.
Regarding the ontogenetic view of alternation of generations, W H Lang (1909) was the first to
work on this aspect. Lang opined that “the differences in the two generations are the result of different
environmental conditions on equivalent germ cells. The spore and the fertilized egg, when first formed,
are conceived to be in perfect neutral condition, without any tendency to form either sporophyte or
gametophyte”. Lang further elaborated that in different environmental conditions (i.e. spore in the open
atmosphere and damp soil, and fertilized egg inside the protected environment of archegonium), both
of them start developing differently. One develops into a sporophyte while the other develops into a
gametophyte. Some later workers, however, gave different views on alternation of generations.
242 � Bryophyta
1. Describe various aspects of alterna on of genera ons in bryophytes and limit your answer in
approximately 500 words.
2. What do you mean by alterna on of genera ons?
3. Explain heteromorphic alterna on of genera ons in one sentence only.
4. Bryophytes show _____ alterna on of genera ons.
5. ‘Archegoniatae’ includes plants having _____, and the term is applied to pteridophytes,
gymnosperms and _____.
6. Draw a diagramma c representa on of alterna on of genera ons.
7. Who discovered alterna on of genera ons in Archegoniatae?
8. Explain the terms “an the c” and “homologous” with reference to alterna on of
genera ons.
9. How will you differen ate between apospory and apogamy?
10. Who was the first to discover apogamy?
11. The term ‘apospory’ was coined by whom and when?
12. Write a detailed account of an the c theory.
13. Who was the first to propose an the c theory?
14. In bryophytes, the complex sporophyte evolved from a simple unicellular zygote gradually by
progressive elabora on through about a dozen stages. Explain these various stages.
15. Who was the first to give some ontogene c views on alterna on of genera ons in
bryophytes?
16. What is homologous theory of alterna on of genera ons? Give some evidences which
support this theory.
16
Phylogeny of
Bryophytes: Role of
Genosystematics
CURRENT STATUS OF MOLECULAR STUDIES
ON BRYOPHYTE PHYLOGENY 16.1
Molecular studies of the recent past have made a definite impact on bryophyte phylogeny. Molecular
data have contributed greatly to develop a phylogeny and modern classification of bryophytes. Widely
known and still used traditional systems of classification of bryophytes in many parts of the world were
based mainly on morphological data, and these have been now significantly revised. Scientists like
Jonathan Shaw and Karen Renzaglia (2004) of USA and AV Troitsky and his team of collaborators
(2007) of Moscow, Russia, have made significant contributions on phylogeny and diversification of
bryophytes with particular emphasis on the role of genosystematics. Several results have been obtained
“from nucleotide sequence data of the nuclear DNA internal transcribed spacers ITS1-21 and the trnL-F
region of the chloroplast genome” (Troitsky et al., 2007).
Shaw and Renzaglia (2004) opined that “the bryophytes comprise three phyla of embryophytes,
that are well-established to occupy the first nodes among extant lineages in the land-plant tree of
life. The three bryophyte groups (hornworts, liverworts, mosses) may not form a monophyletic clade,
but they share life-history features including dominant free-living gametophytes and matrotrophic
monosporangiate sporophytes. Because of their unique vegetative and reproductive innovations, and
their critical position in embryophyte phylogeny, studies of bryophytes are crucial to understanding the
evolution of land-plant morphology and genomes”. These workers have focused also on phylogenetic
relationships within each of the three divisions of bryophytes and relates morphological diversity to
new insights about those relationships. Their “multilocus, multigenome studies have been successful
at resolving deep relationships within the mosses and liverworts, whereas single-gene analysis have
advanced understanding of hornwort evolution” (Shaw and Renzaglia, 2004).”
BRYOPHYTES AND GENOSYSTEMATICS STUDIES 16.2

Bryophytes are a group of most simply organised embryophytes (embryo-containing plants, i.e.
bryophytes and vascular plants) having a dominant gametophyte and lacking a developed vascular
244 � Bryophyta
system. The diploid sporophyte is spore-producing and dependent on the autotrophic gametophyte.
Many bryophytes are often pioneers in extreme habitats and played vital biota-forming roles in the land
settlement by plants.
On the basis of existing catalogues and databases, M S Ignatov (2003) of Russian Academy of
Sciences estimated that three main phyla of modern bryophytes include about 100 (hornworts),
5000(liverworts) and 10,000 (mosses) species. According to Meyen (1987), “the time of origin of
bryophytes and their phyla is into well-determined from paleontological data”, mainly due to bad
preservation of these plants in sediments. However, Sanderson (2003) opined that age of fossil spores
of bryophytes “is 440–450 million years, which is in good agreement with the most reliable molecular-
genetic chronology of the origin of land plants (425–490 million ago)”.
Many studies of bryophytes and genosystematics are still not available, and a majority of the studies
have been done only in the last few decades. Some studies have appeared in the second half of the
1990s, and over half a dozen papers, concerning various bryophyte groups appeared in Bryologist in
2000. Some of the recent studies have been done in 2004 by B Goffinet, V Hollowell and R Magill;
and in 2007 by A E Newton, R S Tagney, K S Renzaglia and J G Duckett. As mentioned also earlier
elsewhere, up to “September 2007, there are 337 entries for hornworts, 5517 for liverworts, and 17,412
for mosses in the GeneBank Database” (Troitsky et al. 2007).
MACROSYSTEMATICS AND ORIGIN OF BRYOPHYTES 16.3

Prior to the researches of molecular phylogenetics, bryophytes were usually divided into two classes,
viz. Bryopsida (mosses) and Hepaticopsida (liverworts). Later on, Anthocerotopsida (hornworts) were
separated from thalloid bryophytes or liverworts. In modern classifications, these three groups have
the rank of divisions, namely Bryophyta, Marchantiophyta and Anthocerotophyta (Kenrick and
Crane, 1997). Several hypotheses of origin of bryophytes have already been proposed, and they were
thought to derive from green, brown, or red algae as “an independent lineage of land-plant evolution,
considered as an intermediate group of evolution from Charales to tracheophytes (vascular plants), or
supposesd to be a result of reduced organisation of more advanced embryophytes” (Kenrick and Crane,
1997).
Cladistic analysis of ultrastructure and ontogenesis of male gametes of several bryophytes has
suggested monophyly rather than paraphyly of bryophytes, which are, “together with Selaginella
(lycophytes), sister to other Lycophytes and other vascular plants” according to Garbary et al. (1993).
Hori et al. (1985) and Van de Peer et al. (1990) were amongst the first to apply molecular data (short
sequences of 5 SrRNA) to phenetic analysis of four species of bryophytes that revealed monophyly of
this group. Waters et al. (1992), however, studied and compared longer sequences (18 S and 26 S rRNA)
of eight species of bryophytes and has shown a “paraphyly of bryophytes, with liverworts in the basal
group and mosses and hornworts sister to vascular plants”. Manhart (1994) made phylogenetic analysis
of the chloroplast rbcL gene sequence of six species of bryophytes and concluded the “polyphyly of
both bryophytes and vascular plants”. Later on, Nickrent et al. (2000) made phylogenetic analysis of
several bryophytes and suggested Anthocerotopsida (hornworts) “as a basal group of embryophytes
and moss-liverwort clade as sister to vascular plants”. All these molecular studies of evolution, thus
give conflicting results on the relationships between three groups of bryophytes (liverworts, hornworts
and mosses) and tracheophytes.
Nashiyama et al. (2004) concluded on the monophyly of bryophytes and made phylogenetic trees
on the basis of their studies of “51 gene sequences from whole chloroplast genomes of 20 higher plants
Phylogeny of Bryophytes: Role of Genosystema cs � 245
and algal species”. Goremykin (2005) also established the monophyly of bryophytes with hornworts
as the most primitive ones, on the basis of his studies of chloroplast genomes from 17 plant species,
including one species each of liverworts, hornworts and mosses.
Samigullin et al. (2002) and Troitsky et al. (2007) constructed phylogenetic trees for 38 species
of bryophytes, 7 species of lycophytes and 2 species of algae “from sequences of inner transcribed
spacers of chloroplast rRNA genes: ITS2, 3, and 4 using three different methods.” According to the
phylogenetic reconstruction of these workers, “hornworts are sister to vascular plants, liverworts
comprise a basal group of land plants, whereas mosses occupy an intermediate position”.
Duff et al (2007) made detailed studies published in Vol. 110 of Bryologist. They have mentioned
that “in a phylogenetic tree of 37 hornwort, 6 liverwort and 4 moss species reconstructed from sequence
analysis of rbcL, nad 5, and 18S rRNA genes, hornworts are the most ancient, and mosses and liverworts
form a clade sister to tracheophytes”.
MARCHANTIOPHYTA OR LIVERWORTS 16.4

Earlier bryologists divided liverworts into two major groups dealing with (i) marchantioid or thalloid
liverworts, and (ii) jungermannioid liverworts including simple thalloid and leafy taxa. Workers, like
Frey and Stech (2005), Crandall-Stotter et al. (2005) and Forrest et al. (2006) developed new systems on
the basis of their detailed genosystematic studies. In these studies, the data-sets used for phylogenetic
reconstructions, included sequences from all major genetic compartments of the cell, viz. (i) chloroplast
trnL-F, rps 4, rbcL, atp B, and psb A, (ii) mitochondrial nad5, and (iii) nuclear rRNA genes. As many as
173 liverwort species were used in these studies, and phylogenetic systems were reconstructed. Basic
findings of all these findings are almost the same. A “backbone” tree of liverwort phylogeny (Fig. 16.1)
was constructed by Forrest et al. (2006) published in Bryologist. In this phylogenetic tree, the basal
position was given to Haplomitriaceae (Haplomitrium) and Treubiaceae (Treubia). Basal position of
these two families is also supported by the anatomical studies of these two genera by Renzaglia et al.
(2007).
On the basis of the recent molecular studies, Blasia, which was earlier included under Jungermannioids,
has now been included under marchantioid liverworts. An independent order (Blasiales) has been
assigned to Blasia under an independent subclass Blasiidae of class Marchantiopsida (Article 2.3.1).
On the basis of phylogenetic studies, He-Nygren (2006) suggested a new system of liverwort
classification, outlined already in Chapter 2. In this system, liverworts have been divided into three
classes, viz. Treubiopsida, Marchantiopsida, and Jungermanniopsida.
ANTHOCEROTOPSIDA OR HORNWORTS 16.5

Represented by only about 100 species, Anthocerotopsida are relatively still unexplored. Genosystematic
studies of Anthocerotopsida have been made by Duff et al. (2004, 2007). A new system has been
proposed. A phylogenetic tree has been reconstructed from sequences of genes like rbcL, nad5, and
18SrRNA. According to these studies, Anthoceros, Folioceros and Sphaerosporoceros “form a separate
clade (subclass Anthocerotidae) sister to most other hornworts affiliated to the subclass Notothylatidae.”
They have also proposed that “Notothylas is closer to the widespread genus Phaeoceros, than the latter
is to the second widespread genus Anthoceros”.
246 � Bryophyta
Fig. 16.1 Phylogene c tree of liverworts suggested by Forrest et al. 2006 (L1, L2 and L3 are leafy
liverworts; ST1 and ST2 are simple thalloids, and CT1 are complex thalloids)
Lieosporoceros, a small hornwort from Central America deserves special mention. It has a peculiarity
of low-level RNA editing. Duff et al. (2007) analysed the phylogenetic relationship of this genus and
compared it with other hornworts. “Based on molecular data and effect of character of RNA editing on
conclusions is reported” by these workers.
BRYOPHYTA OR MOSSES 16.6

According to the modern classification proposed by Kenrick and Crane (1997), Bryophyta is now
treated as a division along with two other divisions—Marchantiophyta and Anthocerotophyta.
It includes mosses (Bryopsida). In all moss families and majority of the genera of mosses, various
DNA regions have now been determined. In the latest system incorporating genosystematics data, all
mosses have now been divided into eight classes, viz. Takakiopsida, Sphagnopsida, Andreaeopsida,
Andreaeobryopsida, Oedipodiopsida, Polytrichopsida, Tetraphidopsida and Bryopsida (Goffinet and
Buck, 2004). Five classes occupying basal positions on the phylogenetic trees are Takakiopsida,
Sphagnopsida, Andreaeopsida, Andreaeobryopsida and Oedipodiopsida. Mosses belonging to other

three classes (viz. Polytrichopsida, Tetraphidopsida and Bryopsida) have peristome. It controls the
dispersal of spores from the capsule of the sporophyte.
According to Troitsky et al. (2007), the position and relationship of many taxa has been drastically
changed in the new system as evident from Fig. 16.2. On the basis of the conclusions made by molecular
phylogenetics, these workers have suggested the following.
“Although the general idea on primitiveness of sphagnous and andreaeid mosses has been
confirmed, some disagreement with systems adopted in both centuries are found, concerning, (i) two
or three basal groups, (ii) inter-relation of main groups distinguished for structure of peristome, and
(iii) pleurocarpous mosses, for which the former system turned out to be absolutely inadequate to new
concepts.” The evolutionary relationships amongst various groups of mosses on phylogenetic trees,
reconstructed from analysis of various genes or their sets, are shown in Article 2.3.2 as opined by these
workers (Troitsky et al 2007).
16.6.1 Importance of Peristome Architecture

In mosses, the main groups are distinguished by their peristome architecture. In very primitive groups
of mosses, the peristome is made up of entire cells, whereas more complexly built ones of coalescent
remnants of cell walls, which allow them to execute multivarious hygroscopic movements (Troitsky
et al., 2007). Molecular data have confirmed such a general evolutionary trend.” Molecular data given
by Ignatova and Ignatova (2003), and Goffinet and Buck (2004) suggest that “more primitive is a
double peristome with opposite elements, and both the double alternate and single peristomes are its
derivatives”.
16.6.2 Pleurocarpous Mosses

The mosses which have a stem with many branches, spreading across the ground are called pleurocarpous
mosses. The reproductive organs in these mosses are borne on short side branches. Approximately 50%
of the present-day moss species are pleurocarpous mosses. The mosses which have an upright stem,
with the reproductive organs at the apex are called acrocarpous mosses. Pleurocarpous mosses form
rapidly extensive coverings on trees, rocks, or soils. It is difficult to classify pleurocarpous mosses
because of ecological diversity of their habitats.
On the basis of the analysis of distinct regions of chloroplast genome and also of mitochondrial
genes, Troitsky et al. (2007) concluded that “diversification of pleurocarpous mosses occurred very
rapidly.” On the basis of such studies, Shaw et al. (2003) opined that “any construction of their reliable
supported phylogeny is impossible”. Troitsky et al. (2007) constructed “phylogenetic trees for 218
samples of pleurocarpous mosses reconstructed in their works from the analysis of nuclear ITS 1-2
sequences and trnL-F of chloroplast genome.”
In determining the phylogeny of mosses, “the choice of genome loci used for the analysis plays
an important role as well” (Troitsky et al. 2007). The set of sequences applicable for these aims is,
however, very limited at present. In plant genosystematics, the nuclear genome ITSs are most variable
in bryophytes among popular loci. In bryophytes, these sequences are used most frequently for studies
at the species level, and can also be successfully employed at higher taxonomic level. Troitsky et al.
(2007) opined further that in revealing of phylogenetic interrelations, “not only construction of trees by
sequences, but also analysis of structural genome rearrangements and localisation of introns may play
very important roles”.
248 � Bryophyta
Brotherus (1924-1925) Goffinet and Buck (2004)

Was unknown Takakiales
Sphagnales Sphagnales Basal groups
Ambuchananiales with specific
Andreaeales capsule
Andreaeales
dehiscence
Was unknown Andraeobryales
Oepodiales Primary peristome-less
Polytrichales Polytrichales Peristome formed
Dawsoniales Tetraphidales from whole cells (nematodontous)
Buxbaumiales Buxbaumiales
Intermediate-
Diphysciales type peristome
Fissidentales Timmiales Double peristome;
Dicranales Encalyptales the outer teeth lie
opposite to the inner
Pottiales Funariales ones(diplolepideous-opposite)
Grimmiales Scouleriales
Bryoxophiales
Simple peristome
Funariales Grimmiales with single ring
Archidiales of teeth
Schististegales (haplolepideous)
Dicranales
Pottiales
Tetraphidales Splachnales
Orthotrichales
Eubryales A
Hedwigiales
Double peristome;
Bryales the outer teeth alternate
Isobryales Rhizogoniales with the inner ones
Hookeriales Ptychomniales (diplolepideous-alternate)
Hypnobryales Hookeriales
P
Hypnales
Fig. 16.2 Interrela onships between the systems of mosses suggested by Brotherus (1925) and
Goffinet and Buck (2004) as also illustrated by Troitsky et al. (2007) (A, Apocarpous
mosses; P, Pleurocarpous mosses)
MAJOR ROLES OF MOLECULAR PHYLOGENETICS 16.7

In solving several taxonomic problems, particularly in bryophytes, molecular phylogenetics has several
major roles to play. Of these, some are listed below:
1. It has “introduced a determining contribution to the solution of phylogenetic and systematic
problems in bryophytes”.
2. It “opened wide perspectives for populational and micro-evolutional studies” in bryophytes.
3. Molecular phylogenetics “has provided strong impulse to the search for new morphological
and anatomical markers of phylogenesis” and to provide new concepts of “morphological
evolution pathways”.
4. It has revealed “key groups in evolution”, which has “put forward new tasks for comparative
anatomy.”
5. Molecular phylogenetics has a definite role in determining “the tasks for investigations
on evolution of distinct systems: genetic, biochemical, physiological, anatomical and
morphological”.
1. Write an essay on phylogeny of bryophytes with par cular reference to role of

genosystema cs.
2. The evolu onary history of an organism or taxonomic group of organisms is known as _____
3. _____ is the gene c material on the sets of chromosomes in a cell.
4. Embryophytes are embryo-containing plants, i.e. all vascular plants and_____.
5. Major studies on bryophytes and genosystema cs have been done during last ____
decades.
6. Give an account of macrosystema cs and origin of bryophytes.
7. Hornworts include members of the class _____
8. A method of classifica on in which the rela onships between organisms are represented by
a diagram, or cladogram, based on selected shared characteris cs is known as _____
9. Make a phylogene c tree of liverworts as suggested by Forrest et al. (2006).
10. Genosystema c studies of which of the following are s ll least explored?
(a) Marchan ophyta, (b) Anthocerotopsida, (c) Bryophyta
11. According to modern classifica on, Bryophyta is treated as a division along with two more
divisions, namely Anthocerotophyta and _____.
12. Differen ate between pleurocarpous mosses and acrocarpous mosses in about 100 words.
13. Enumerate some major roles of molecular phylogene cs.
17
Morphogenesis
WHAT IS MORPHOGENESIS? 17.1
Morphogenesis is “the development of shape and structure of organs and tissues”. Or, it may also be
defined as “the developmental changes that give rise to the adult form from the zygote”.
LIMITS OF MORPHOGENETIC STUDIES IN BRYOPHYTES 17.2

Morphogenesis is such a vast subject that it is not possible to specify its limits in one chapter of a book
of this nature. The studies made on the subject during the last fifty years are so enormous that even a
brief mention of all of them is neither possible nor within the scope of this book. Only certain selected
undermentioned aspects are listed and discussed, in general, here briefly:
1. Germination of spore
2. Initiation of bud and its growth into gametophyte
3. Physiology of sex expression
4. Growth of diploid sporophyte
5. Vegetative propagation
6. Metabolism
7. Senescence
8. Physiology of rhizoid formation
GERMINATION OF SPORE 17.3

Stephen (1928), Mohr (1956) and many others observed that there is an absolute requirement of light
for spore germination. But, workers like Kessler (1914), Meyer (1948) and many others reported that
spores of some mosses also germinate in darkness on inorganic media. Besides light, some other
factors responsible for germination of spores in bryophytes are (i) temperature (optimum temperature
16–25°C), (ii) moisture (in the form of liquid water or large amount of air humidity), (iii) pH (optimum
Morphogenesis � 251
pH is more on the acid side but sometimes it is neutral or slightly on the alkaline side), and (iv) more
amount of oxygen, etc.
Several studies have confirmed that sucrose accelerates the germination of spores in bryophytes
(Benson-Evans, 1953; Hoffman, 1964; Valanne 1966).
Growth substances (e.g. IAA) have a marked effect on the polarity of spores. Gibberellic acid and
kinetin induce germination in darkness. Vaarama and Taran (1963) observed that there was a clear
promotion of germination by gibberellic acid. Valanne(1966) reported that low concentrations of
gibberellin have been found to promote germination in Ceratodon purpureus.
17.3.1 Liverworts
In Hepaticopsida, the spores do not germinate in dark, even if sugar is present, and according to Inoue
(1960), blue light plays an important role in spore germination in many taxa. Mohr (1963) observed
that spore germination can be induced by both blue and far-red radiations in Sphaerocarpos donnellii.
Steiner (1964) studied the spore-germination aspect further and observed that it can be controlled
by phytochrome as well as high-energy reaction, and while functioning, there exists an interaction
between phytochrome and high-energy reaction.
It has also been established in some liverworts that spore germination in blue and green light can
be promoted if there is a simultaneous irradiation of these two lights. On the other hand, germination
inhibits in far-red light. Regarding spore germination and quality of light, Steiner (1964) also proved
that red light is more effective at the beginning of day and far-red at the end of the day. Doyle (1963)
observed that red, far-red, green and blue lights are equally effective in promoting spore germination
in Sphaerocarpos cristatus.
17.3.2 Mosses
In mosses, spore germination passes through two phases: (i) the first phase results into an increase
in volume due to absorption and uptake of water, and (ii) the second phase starts by rupturing of the
exospore and intensive greening of plastids due to light. Rhizoid initiation also takes place in the
second phase.
It has also been proved that requirement of light in the germination of spores in mosses is associated
with the phytochrome system. Valanne (1966) detected phytochrome system in Ceratodon purpureus
and a few more mosses, while Egunyomi (1979) detected it in Octoblepharum albidum. The phytochrome
system has, however, not been detected in Funaria hygrometrica. According to Krupa (1967), blue and
far-red lights prove quite effective in inducing spore germination in Funaria hygrometrica. Different
studies suggest that mosses, in general, differ in their response to different spectra of light. Red light
(640 to 680 nm) is more effective in including spore germination in Physcomitrella patens. It has also
been proved by Kass and Paolillo (1977) that in Polytrichum, the chloroplasts of germinating spores
replicate more in light than in darkness.
252 � Bryophyta
INITIATION OF BUD AND ITS GROWTH INTO

GAMETOPHYTE 17.4
17.4.1 Condi ons Determining Bud Ini a on
The major conditions which determine the initiation of buds, specially in mosses, include (i) effect of
light, (ii) effect of temperature, (iii) effect of carbohydrates, and (iv) effect of growth regulators.
Effect of light on initiation of buds in mosses has been studied by many different workers, and it has
been finally established that moss protonema, when reared under high light intensities, readily forms
buds. Besides intensity, the quality of light is also important in bud initiation.
Effect of temperature on bud initiation has also been studied in several mosses including Funaria
hygrometrica, Phascum cuspidatum and Physcomitrium turbinatum. It has been established that a
majority of mosses have a certain temperature range during which the buds are formed.
Regarding the effect of carbohydrates, it can be generalised that an attainment of a certain minimum
concentration of sugar in the moss protonema is a necessary prerequisite for initiation of buds.
Growth regulators (IAA, GA, kinetin, etc.) have a definite effect on formation of buds in mosses.
Higher concentrations of auxins have a marked inhibitory effect on the formation of shoot buds on the
protonema. Kinetin very strongly stimulates the process of bud initiation in mosses.
17.4.2 Growth of Gametophyte of Some Liverworts, Hornworts and Mosses

Consult Chapter 13 (Articles 13.5, 13.6 and 13.7).
PHYSIOLOGY OF SEX EXPRESSION1 17.5

Sex in several bryophytes is determined genotypically, but in many others, it is determined phenotypically,
i.e. under environmental control. Wettstein (1924) believed that phenotypic determination of sex
in mosses is an interesting process. Androgynous mosses, which are protrandrous, show “tissue
differentiation as male and female organs have bisexual potentialities, i.e. plants regenerated from this
tissue again produce both sexes in a genetically determined sequence”. Several mosses show genotypic
sex expression, e.g. Bryum argenteum. In such cases, some spores give rise to female gametophytes
and others to male gametophytes. Differing from this, the diploid gametophytes, regenerated from the
sporogonium, possess both male and female sex organs.
In Sphaerocarpos donnellii, out of the four spores produced by a tetrad, two produce male and two
the female gametophytes. The females have a very large chromosome, perhaps X-chromosome, and
the males have a much smaller homologue, perhaps Y-chromosome. Similar pairs of sex chromosomes
are present in many other dioecious liverworts and mosses.
It was observed by Bauer (1959) that “diploid gametophytes, regenerated from the sporogonium of
dioecious Splachnum luteum, formed archegonia but failed to form antheridia if they were transferred
at frequent intervals.” Three main points are observed in the stabilisation of the male sexual tendency
in Splachnaceae, according to Bauer (1961, 1963).
1
Also refer Chapter 14.
(i) “Protonema, regenerated from the sporogonium, is at first potentially male;

(ii) Sex-expression determination is at first liable for the effect can be reversed by rejuvenation,
and
(iii) After the male sex organs attain maturity, the maleness of the strain becomes stabilised”. Bauer
(1963) also studied and confirmed that in Splachnum rubrum, “the male sexual characters
appear on a genotypically determined female plant”. Lorbeer (1936) and Heitz (1942) are
some of the workers who opined that in Hepaticopsida, the transformation of genotypically
determined females has been successfully obtained several times by giving X-ray treatment,
i.e. by means of mutation. In Splachnum ampulaceum, S. luteum and S. rubrum, the male
plants are formed from the females in the same fashion irrespective of whether they are of
phenotypic or genotypic sex determination.
GROWTH OF DIPLOID SPOROPHYTE 17.6

Light plays a definite role in the growth and expansion of sporophyte in mosses. In Funaria hygrometrica,
it was observed as early as 1886 by Haberlandt that light is required for proper development of its
capsules. The spore sac failed to expand in darkness in Funaria even if the gametophyte and seta were
exposed to light. Paolillo and Bazaz (1968) also observed the role of light in controlling expansion of
capsule in Polytrichum and opined that “it appears to be morphogenetic rather than photosynthetic”. In
capsule expansion, the red light is more effective than white, blue, or green light of equal energy. Krisko
and Paolillo (1972) opined that in Polytrichum, comparatively low irradiations are needed for expansion
of capsules, and French and Paolillo (1976) also proved the same in Funaria hygrometrica.
The phenomena of apogamy (asexual reproduction in which embryos and propagules are produced
without meiosis occurring) and apospory (production of a diploid gametophyte from vegetative cells
of the sporophyte, that is, without the production of spores) are also of importance from evolutionary
and morphogenetic point of views. For more details of these two phenomena, consult Chapter 20.
VEGETATIVE PROPAGATION2 17.7

17.7.1 Hepa copsida (Liverworts)
An important role is played by light in the vegetative propagation by promoting the formation of
gemmae and other types of propagules in liverworts. Gemma is a small group of cells in the form of
a bud that will give rise to a new individual. Gemmae are produced in cup-shaped structures on the
surface of thalloid liverworts.
In liverworts, the vegetative propagation through gemmae is a light-controlled response. It is so
because gemmae do not germinate in dark. In Marchantia polymorpha, the rhizoid formation and
further growth of gemmae is controlled by the phytochrome system. According to Otto and Halbsguth
(1976), red light is essential for the rhizoid formation on gemmae in M. polymorpha. Kaul and Kaul
(1974) observed that germination of gemmae in Marchantia nepalensis takes place only in red, orange
2
Also refer Chapter 18.
254 � Bryophyta
or green light. Also in Lunularia cruciata, the formation of rhizoids on gemma is very sensitively
controlled by red or far-red light system. If given for a very short period, the far-red light inhibits
rhizoid formation completely in L. cruciata.
17.7.2 Bryopsida (Mosses)

Phytochrome is a pigment which controls many of the physiological responses of plants to light. This
actually absorbs red and far-red wavelenghts of light. In Mnium affine, regeneration of protonema from
the cells of its isolated leaves is under the control for its phytochrome-mediated system. Giles and von
Moltzahn (1967) observed that action of blue light is similar to that of red light in M. affine, and this
action is reversible by far-red light. Both these researchers also isolated phytochrome from M. hornum
and M. undulatum. Demkiv (1971) studied regeneration in cells of protonema in Funaria hygrometrica
and concluded that it occurs in red and blue light, and is probably influenced by phytochrome. In
Aulacomnium palustre and Tetraphis pellucida, the germination of propagules is controlled by
phytochrome.
Kumra (1977) reported that red light promotes maximum gemmae production in Bryum klinggraeffii,
and blue light and green light are next in order of effectiveness. In yet another study, Jenkins and
Cove (1983) observed the light requirements for regeneration of protoplasts of Physcomitrella patens.
A simple regeneration sequence is followed by this moss, which include (i) synthesis of cell wall,
(ii) formation of an asymmetric wall, (iii) division of this asymmetric wall, and (iv) further extension
and division to produce a new chloronemal filament. Except that of cell-wall formation, all these
processes require light for their completion. For cell division, red light is more effective than blue or
far-red light.
METABOLISM 17.8
The sum of the chemical reactions which occur in an organism or a cell is known as metabolism. It
involves the breakdown of organic compounds, and releasing energy that is used in the synthesis of
other compounds. In bryophytes, it is the quality of light that controls and affects, directly of indirectly,
the growth and development, which actually take place by synthesis of many endogenous morpho-
regulatory substances.
Several experiments have so far been performed on metabolism in different members of bryophytes.
A few such examples are mentioned here:
1. In the vegetative thalli of Marchantia polymorpha, Melstrom et al. (1974) detected three
endogenous gibberellin-like substances. If the photoperiod is increased from 12 to 18 hours, it
resulted in the quantitative difference in the activity of these gibberellin-like substances, and
the result is observed in the form of increase in growth and elongation of thallus.
2. A compound (lunularic acid), which controls growth and drought resistance, has been isolated
from Lunularia cruciata. Growth of gemmae, while they are still within gemma cups, is
checked by lunularic acid.
3. Phytochrome regulates the synthesis of acetylcholine in the moss callus, regenerated from the
seta of hybrid sporophyte of Funaria hygrometrica and Physcomitrium pyreforme. In these
hybrid sporophytes, red light promotes synthesis of acetylcholine while far-red inhibits its
synthesis.
4. In Ceratodon purpureus, light induces the synthesis of a cellular division factor by the cells of
its protonema.
5. According to Chopra and Kumra (1978), a morphoregulatory substance, produced by the
protonema of Bryum klingraeffii, controls the formation of gemma in this moss.
SENESCENCE 17.9
The process of growing old before death is known as senescence. The quality of light controls senescence
in Marchantia polymorpha. It has been shown in this liverwort that its thalli remain green if a daily one
hour photoperiod of white light is given. Its tissues, however, start showing bleaching symptoms when
daily one-hour photoperiod is terminated with a brief irradiation of far-red light. While bleaching its
chlorophyll is lost and there is simultaneous breakdown of its cell organelles and cytoplasm. There is a
clear implication of phytochrome in the control of senescence by light.
PHYSIOLOGY OF RHIZOID FORMATION 17.10

Experiments of LaRue (1942) and others have proved that auxins stimulate the rhizoid production in
bryophytes. Inhibition of rhizoid production induced by darkness in gemmae of Marchantia polymorpha
was overcome by applying IAA at a concentration of 0.1–0.01 ppm. It was shown by LaRue (1942)
that rhizoid formation gets stimulated by applying IAA and IBA in lanolin on leaves and sporophytes
of several mosses and also on thalli of Conocephalum conicum.
The effect of indole-acetic-acid on the regenerative behaviour of the isolated tissue of the rhizoids
of Riella helicophylla was studied by Stange (1958). Initially, a majority of the cells underwent
regenerative cell division. IAA did not cause rhizoid formation after isolation of individual cells or a
small group of cells by plasmolysis. Stange (1958) observed that IAA influences or promotes rhizoid
production in Riella helicophylla.
Morphogenetic responses of several growth substances (e.g. IAA, IBA, 2, 4-D, etc.) on the thallus
of Marchantia was studied by Kaul et al. (1962). They observed that if a concentration of 1 ppm or
more of growth substances is applied, it stimulated rhizoid formation, but thallus growth was inhibited.
Major rhizoid-inducing growth substances according to these workers are NOA (Naphthalene
Oxyacetic Acid), NAA, (Napthalene Acetic Acid) and TCPA (Trichlorophenoxy Acetic Acid). They
induced rhizoid formation not only on ventral but also on dorsal surface of the thalli of Marchantia. 0.1
ppm of NAA stimulates the production of rhizoids but not of the thallus. According to Allsopp et al.
(1968) also, auxins have a pronounced effect on rhizoid formation in Marchantia polymorpha. They
(IAA, NAA and 2, 4-D) not only induced production of an increased number of rhizoids but also their
formation from both upper and lower surfaces of the gemma in this liverwort.
256 � Bryophyta
1. Give an overview of morphogenesis in bryophytes in about 1000 words.

2. Define the following terms:
(a) Morphogenesis (b) Apogamy
(c) Apospory (d) Phytochrome
(e) Metabolism (f) Senescence
3. Explain germina on of spore in bryophytes, keeping in mind the phenomenon of
morphogenesis.
4. What are the major condi ons which determine ini a on of bud in mosses?
5. Describe physiology of sex expression in bryophytes in about 250 words.
6. Is it true that light plays a definite role in the growth and expansion of sporophyte in
mosses?
7. The pigment which controls many of the physiological responses of plants to light is named
as _____.
8. The sum of chemical reac ons, which occur in an organism or a cell, is known as _____.
9. The process of growing old before death is known as _____.
10. Elaborate the abbrevia ons NAA and TCPA.
18
Vegetative
Reproduction and
Regeneration in
Bryophytes
DIFFERENCE BETWEEN VEGETATIVE REPRODUCTION
AND REGENERATION 18.1
Vegetative reproduction may be defined as a type of asexual reproduction in which a whole new plant
is produced from an organ, e.g. a tuber, which is not involved in sexual reproduction. In the absence of
mutation, the offspring of the vegetative reproduction will be genetically identical to the parent plant.
Regeneration is (i) the growth of new tissue on a part of a plant that has been damaged; or (ii) the
growth of new plants from perennating organs, e.g. rhizome. It may also be defined as regrowth of
tissues or organs; or the formation of new plants from cultured tissues.
METHODS OF VEGETATIVE REPRODUCTION

IN BRYOPHYTES 18.2
Bryophytes reproduce vegetatively by various means. In several of their dioecious species, the plants
reproduce mainly by vegetative methods, and some such species have even ceased to reproduce sexually.
C Correns (1899), a famous bryologist, was the first to summarise various methods of vegetative
reproduction in bryophytes with main emphasis on mosses (Bryopsida), while F Cavers (1903) was
the first to give an authentic and detailed account of various methods of vegetative reproduction in
liverworts (Hepaticopsida) in his two articles “on asexual reproduction and regeneration in Hepaticae”,
published in the journal New Phytologist. The detailed studies of both these bryologists on vegetative
reproduction in bryophytes have been beautifully summarised by the well-known Indian author N S
Parihar (1987), and an outline of the same is presented here in this account.
258 � Bryophyta
1. Death and Decay of Older Parts In a majority of Hepaticopsida and Anthocerotopsida,

the younger branches of dichotomously dividing thallus get isolated due to progressive death and
decay of the older posterior parts. Such isolated parts start functioning as new thalli (Fig. 18.1 A-C) on
return of favourable conditions. This is the most common method of vegetative reproduction in thalloid
bryophytes, particularly in liverworts and hornworts. This is also seen in those mosses (Bryopsida)
which have prostrate rhizomes bearing erect branches.
Fig. 18.1 A-C Isola on of younger dichotomies due to death and decay
2. Adven ous Branches The term ‘adventitious’ is applied to a plant part developed out of the
usual order or in an unusual position. In thalloid bryophytes, adventitious branches usually develop from
the underside of the midrib. Upon separation from the parent plant, the adventitious branch develops
into a new plant, e.g. Anthoceros laevis, Asterella, Blasia, Corsinia, Dumortiera, Marchantia, Pellia,
Reboulia, Riccia fluitans, Sphaerocarpos, Targionia, etc.
3. Innova ons A young offshoot from the stem is known as innovation. On being separated,
and falling on the suitable substratum, the innovations develop into new plants, e.g. Sphagnum and
many acrogynous Jungermanniales. In Sphagnum, the innovation grows more vigorously than the
other branches, continues its upward growth, and takes all the characteristics of the main axis. It is an
effective method of multiplication in Sphagnum.
4. Leaf Cladia A small detachable branch originating from the individual cells of the leaf is known
as leaf cladia. On being detached, leaf cladia develop into new plants, e.g. Frullania fragilifolia,
Plagiochila, etc.
5. Stem Cladia It is a small detachable branch which originates from the individual cells of the stem.
It occupies almost the same position on the stem as the sex-organ bearing branch. Stem cladia develop
on the stem in several leafy liverworts and mosses, e.g. Bryopteris fruticulosa, Drepanolejeunea, etc.
6. Whole Shoots All and complete shoots get separated from the parent plant, and on getting
suitable water and other requirements develop into new plants, e.g. Pohlia nutans.
7. Shoot Tips In Campylopus, Polytrichium and some other mosses, tips of the shoots get separated
and develop into new plants.
8. Modified Branch of Budlike Form In some species of Bryum and many species of Pohlia
and some more mosses, the organ shed is a modified branch of bud-like form. This type of organ is
Vegeta ve Reproduc on and Regenera on in Bryophytes � 259
strictly a deciduous branchlet and quite capable of developing into a new plant. In some taxonomic
works of bryophytes, such organs have also been termed as bulbil or gemma.
9. Tubers Technically, a tuber is a “thick underground stem in which food is stored”, or, “a
swollen underground stem acting as a storage and perennating organ”. Some define tuber as a “swollen
part of a stem or root, usually modified for storage”. In bryophytes, tubers develop in several species
of liverworts as well as mosses. Some of the common tuber-forming species are Riccia billardieri,
R. discolor, Geothallus tuberosus, Asterella angusta, Aitchisoniella himalayensis, Conocephalum
conicum, Fossombronia tuberifera, Sewardiella tuberifera, Anthoceros himalayensis and A. laevis
(Fig. 18.2 A-B).
Tubers
A B
Fig. 18.2 A-B Tubers on the thallus of Anthoceros himalayensis (A) and. A. laevis (B)
10. Gemmae Gemmae (singular: gemma) are “small groups of green cells, produced in cup-shaped
structures on the surface of thalloid liverworts.” Gemmae are usually dispersed by splashes of rain.
Some define gemmae as “a bud that will give rise to a new individual, e.g. the multicellular structure
in algae, pteridophytes and specially bryophytes”. These are the means of vegetative propagation and
form abundantly in liverworts (Hepaticopsida), to some extent in hornworts (Anthocerotopsida) and
to a lesser extent in mosses (Bryopsida). Gemmae are not formed in Sphagnales. Gemmae reported in
different groups of bryophytes are listed below.
(a) Gemmae of Liverworts (Hepa copsida) Many types of gemmae are produced in Hepaticopsida.
Of these, some are listed below:
(i) One to three-celled gemmae developing on the stem apex, e.g. Cephalozia bicuspidata,
Lophozia heterocolpa.
(ii) One- to three-celled gemmae developing on the leaves, e.g. Cephalozia francisci, Lophozia
barbata.
(iii) Two-celled endogenous gemmae developing within any external cell of the thallus, e.g.
Riccardia multifida (Fig. 18.3A), Haplozia caespiticia.
(iv) Three- to four-celled gemmae developing in the axils of the leaves, e.g. Treubia.
(v) Stalked, multicellular, discoid gemmae formed on the dorsal surface of the thallus inside
gemma cups, e.g. Marchantia (Fig. 18.3 B), Lunularia.
260 � Bryophyta
Gemmae cup
Gemmae
Gemma
Gemmae
Thallus
Gemma
A B C D
Fig. 18.3A-D Gemmae of some bryophytes. A, Riccardia mul fida; B, Ver cal sec on of the thallus of
Marchan a polymorpha through gemma cup; C, Radula complanata; D, Metzgeria uncigera
(vi) Subspherical gemmae produced in large number in flask-shaped gemma receptacles e.g.
Blasia.
(vii) Star-shaped gemmae developing on the dorsal surface of the thallus, e.g. some species of
Blasia.
(viii) Discoid, multicellular gemmae developing on leaves, e.g. Rudula complanata (Fig. 18.3C),
Leptocolea.
(ix) Discoid multicellular gemmae developing on erect gemmiferous branches, e.g. Metzgeria
uncigera (Fig. 18.3D).
(b) Gemmae of Hornworts (Anthocerotopsida) Some Anthoceros species bear gemmae. In A.

glandulosus, several gemmae develop along the margin of the thallus, and a few also develop on the
dorsal surface. Gemmae have also been reported in some more species, such as A. formosae and A.
propaguliferus.
(c) Gemmae of Mosses (Bryopsida) Six types of gemmae developing on different plant parts of
mosses are listed as under:
(i) On Rhizoids of Leafy Shoots These are the multicellular gemmae developing on the rhizoids
of leafy shoots, e.g. Barbula convoluta, Bryum erythrocarpum, etc.
(ii) At the Base of Stem In Bryum erythrocarpum (Fig. 18.4A) and B. rubens, multicellular and
globular gemmae develop at the base of the stem.
(iii) At the End of Leafless Stalks In Aulacomnium androgynum (Fig. 18.4 B), stalked fusiform
gemmae develop at the end of leafless stalks.
(iv) At the Tip of the Shoot In Tetraphis pellucida (Fig. 18.4 C), green, stalked, multicellular and
lenticular gemmae develop at the tip of the shoot. Widened leaves form a cup-like structure
around such gemmae.
(v) On the Stem and Branches In Pterygynandrum filiforme, smooth, golden-brown gemmae
develop on the stems and branches. Such gemmae are ovoid and stalked bodies. These are
bicelled structures and develop on colourless stalk made of three or more cells.
(vi) On the Leaves In Torula papillosa, Ulota phyllantha (Fig. 18.4D) and some more bryophytes,
multicellular, articulate gemmae develop on the leaves.
Fig. 18.4 Gemmae found in some bryopsida. A, Bryum erythrocarpum; B, Aulacomnium

androgynum; C, Tetraphis pellucida; D, Ulota phyllantha
11. Primary Protonema In almost all mosses, the spore germinates into a young filamentous
and branched gametophyte, known as primary protonema. It usually breaks into smaller parts
or fragments, and each such fragment is capable of developing into a new protonema and forms
gametophyte initials.
12. Secondary Protonema A protonema-like structure, developing by other methods than

from the germination of spores is known as secondary protonema. It is also a means of vegetative
reproduction in Bryopsida (e.g. Funaria hygrometrica). Secondary protonema develops (i) either from
the rhizoids of a leafy gametophore when exposed to light, or (ii) from almost any separated living
part of the moss plant such as stem, leaves, sex organs, or even paraphysis and sterile cells of seta and
capsule.
BOTANICAL NOTE ON REGENERATION IN BRYOPHYTES 18.3

As mentioned earlier also, regeneration is technically “the growth of new tissue on a part of a plant
that has been damaged, or it is the growth of new plants from the perennating organs, e.g. rhizomes”.
Fulford (1956) defined “regeneration” as a process in which a new plant is formed from a presumably
adult cell which has undergone differentiation back to an activated condition.” Regeneration actually
means the capacity of an organ, tissue or cell to give rise to the entire organism. According to Wilmot-
Dear (1980), regeneration of differentiated cells follows injury or isolation from the meristem, and the
isolation may be physical or physiological.
Generally, bryophytes possess enormous potentialities for regeneration. Regeneration of the
physiologically isolated parts of both sporophytic and gametophytic generations takes place readily in
an artificial medium. Regeneration in nature may be caused because of the attacks of fungi, insects, from
mechanical injuries of many sorts or even from unfavourable conditions which impede the function of
plant parts. According to Stange(1964), regeneration does not include “normal physiological processes
in which new growth is initiated as normal vegetative reproduction or formation of secondary meristems
or meristemoids in plants”.
262 � Bryophyta
Liverworts regenerate with particular readiness among bryophytes. For example, in Sphaerocarpos,
regeneration occurs from single cells or a group of adjacent cells from almost anywhere on the thallus.
They first form a globular, cylindrical or ribbon-like body, which soon develops into a new typical
thallus, as is formed from a germinating spore of Sphaerocarpos. Among Jungermanniales (e.g.
Fossombronia) also, plantlets develop frequently from single cells in the leaves and their development
is almost in the similar fashion as that of a spore. Mehra and Pahwa (1971) observed in thalli of
Fossombronia himalayensis that they regenerate from rhizoids of starved cultures.
Many Bryopsida (mosses) also have excellent power of regeneration. Several studies have been made
in India by many different workers, including Kachroo (1954) on Physcomitrium pyreforme, Chopra and
Sharma (1956) on Pogonatum, and Banerji and Sen (1957) on Barbula indica. Different mosses show
different regenerative capacity from many of the body parts such as leaves, antheridia, archegonia or
as protonemal outgrowths from different parts of the sporophyte. In mosses, the regeneration from the
leaves occurs only if they are detached from the axis, as shown by many workers including Gemmell
(1953) in Atrichum and Meyer (1942) in Physcomitrium turbinatum. Several Japanese workers have
made extensive contribution on the regeneration of detached leaves in mosses, and, in general, have
opined that “the manner of regeneration of leaves is similar to that of the germination of spores in
mosses” (Noguchi and Mizuno, 1959). Chopra and Kumar (1961) generalised that “the percentage of
regeneration increases with the advance of age in different species of Atrichum”, a moss. They have
shown that “lower percentage of regeneration of the younger organs is due to scanty food reserves as
they are actively used by these organs in their growth and other activities”.
Chopra and Kumra (1988) have compiled a list of “some specific examples of regeneration from
organs other than thallus, protonema, stem, leaf, and seta”. Some of these specific examples are listed
below:
1. Antheridial stalk—Bryum cellulare (Narayanaswami and Lal, 1957)
2. Archegonial neck—Mnium (Wettstein, 1924)
3. Archegonial stalk—Rhodobryum (Narayanaswami and Lal, 1957)
4. Archegonial venter – Funaria (Correns, 1899)
5. Calyptra – Barbula indica (Narayanaswami and Lal, 1957; Fig. 18.5 A)
6. Jacket of capsule—Funaria (Kumra and Chopra, 1980)
7. Perianth—Fossombronia (Mehra, 1976)
8. Perichaetial leaves—Octoblepharum (Egunyomi et al., 1980)
9. Rhizoids—Physcomitrium coorgense (Narayanaswami and Lal, 1957)
10. Vaginula—Barbula indica (Fig. 18.5B; Narayanaswami and Lal, 1957)
Major factors which affect regeneration are light, radiation, pH, humidity, season, temperature,
reserve food material, wounding, location of the plant, size of the fragment and age.
For more details of regeneration in bryophytes, readers may consult Biology of Bryophytes by
Chopra and Kumra (1988).
Fig. 18.5 Barbula indica showing regenera on from calyptra (A) and vaginula (B)
1. How can you define the two terms? (i) Vegeta ve reproduc on, (ii) Regenera on.
2. Write an essay on various methods of vegeta ve reproduc on in bryophytes.
3. A thick underground stem in which food is stored is known as _____.
4. Is the word “gemmae” singular or plural?
5. Give a detailed account of gemmae in bryophytes.
6. “Gemmae are formed in all the three major groups of bryophytes”. Comment.
7. Give a brief illustrated account of gemmae of Hepa copsida.
8. How do mosses reproduce by protonema?
9. Write a brief scien fic note on regenera on in bryophytes in about 200 words.
10. Define regenera on.
11. Besides thallus, protonema, stem and leaves, mosses exhibit the phenomenon of regenera on
also from almost all parts of their body. Jus fy giving suitable examples.
12. Innumerate major factors which affect regenera on in bryophytes.
19
Origin and Fate of
Archesporium in
Bryophytes
WHAT IS ARCHESPORIUM? 19.1
The tissue that gives rise to spore mother cells is known as archesporium. The archesporium is
actually the first cell generation of the sporogenous tissue. In almost all bryophytes, it appears as a
continuous tract of tissue, which occupies (i) usually a central position, as in majority of liverworts
(Hepaticopsida), or (ii) a more or less superficial position between the wall and the columella, as
in majority of hornworts (Anthocerotopsida) and mosses (Bryopsida). The cells of the archesporium
divide and redivide several times to form a large number of sporogenous cells.
PRODUCTS OF ARCHESPORIUM 19.2

Sporogenous tissue, produced from the archesporium, develops into several types of cells in different
bryophytic genera, as shown below:
1. Sporocytes These are produced in all bryophytes and are also called spore mother cells. They
are diploid in nature, divide by meiosis and produce haploid spores.
2. Abor ve Nurse Cells These are produced along with sporocytes in some species of Riccia.
They abort soon and form a nutritive fluid for the developing spore mother cells.
3. Persistent Nurse Cells In the sporogonium of some genera, such as Geothallus, Riella and
Sphaerocarpos, some cells persist for quite some time along with spore mother cells. These are called
persistent nurse cells.
4. Elaters Elaters are a bunch of long, thin cells in the capsule of the sporophyte of several
liverworts, e.g. many members of Marchantiales (e.g. Marchantia) and Jungermanniales. Elaters have
spiral thickenings of the cell wall. They alter their position with changes in humidity, and help in the
dispersal of spores from the capsule.
Origin and Fate of Archesporium in Bryophytes � 265
5. Apical Cap of Sterile Cells These are the cells present in the form of a cap on the apical
side of the capsule of the sporophyte of some bryophytic genera, as in some Marchantiales (e.g.
Marchantia).
6. Elaterophore In some Jungermanniales (e.g. Pellia, Riccardia), some large-sized sterile cells
are present in the capsule either at the base (e.g. Pellia, Fig. 19.1 A) or at the top (e.g. Riccardia, Fig.
19.1 B). These cells develop spiral thickenings on their walls and form a somewhat compact structure
called elaterophore. Some elaters also get attached on this, making it a thick and more distinct structure
inside the capsule.
7. Pseudoelaters In Anthocerotopsida (e.g. Anthoceros, Fig. 8.7 M) the sporogenous tissue gives
rise to sporocytes and some elaters or pseudoelaters.
Fig. 19.1 Elaterophore at the base of capsule in Pellia epiphyla (A), and at the top
of the capsule in Riccardia pinguis (B)
266 � Bryophyta
FUNDAMENTAL EMBRYONIC LAYERS OF CAPSULE 19.3

The amphithecium and endothecium are the two fundamental embryonic layers of capsule in
bryophytes. Usually, the diploid zygote divides and redivides to form a multicellular, more or less
spherical mass of 20 to 40 cells. They divide by a periclinal division to form an outer layer, called
amphithecium and an inner central mass of cells, called endothecium (e.g. Riccia). The archesporium
(the first cell generation of sporogenous tissue) is amphithecial in origin in some bryophytes (e.g.
Anthoceros), while in others it is endothecial in origin (e.g. Marchantia).
ORIGIN, POSITION AND FATE OF ARCHESPORIUM

IN SOME SELECTED GENERA 19.4
The origin, position and fate of the archesporium in majority of the genera of Hepaticopsida,
Anthocerotopsida and Bryopsida, discussed earlier in Chapters 4 to 10, have already been described. A
brief summary of some of the selected genera is given in Table 19.1.
Table 19.1 Origin, posi on and fate of archesporium in some common genera of bryophytes
NO. NAME OF GENUS ORIGIN POSITION FATE

1. Riccia Endothecial; entire Centrally-located; forms Almost all sporogenous
endothecium functions as entire sporophyte except cells start functioning as
archesporium and forms single-layered wall. spore mother cells, which
sporogenous tissue. divide reductionally to form
haploid spores; a few pe-
ripheral cells, called nurse
cells, however, degenerate
and form nutritive fluid.
2. Marchantia Endothecial; entire Entire cavity of capsule Half of the sporogenous
endothecium functions remains filled with sporog- tissue forms spore mother
as archesporium, divides enous tissue surrounded by cells and remaining half
and redivides several single-layered wall of the forms elaters. Some cells
times and forms sporog- capsule. at the top remain sterile and
enous tissue. form the apical cap.
3. Porella Endothecial; entire Entire cavity of capsule Entire sporogenous tissue
endothecium functions remains filled with sporog- forms spore mother cells
as archesporium, which enous tissue, which remains and elaters. Elaterophore is
divides and redivides to surrounded by two or more- absent.
form sporogenous tissue. layered wall of the capsule.
4. Pellia Same as in Porella. Same as in Porella. Some sporogenous tissue
forms basal elaterophore
(Fig. 19.1A); remaining
sporogenous tissue forms
spore mother cells and
elaters.
Contd.
Origin and Fate of Archesporium in Bryophytes � 267
5. Riccardia Same as in Pellia and Same as in Pellia and Same as in Pellia except that
Porella. Porella. the elaterophore is at the top
of the capsule (Fig. 19.1 B).
6. Anthoceros Amphithecial; arches- Located more superficially Regarding the fate of the
porium originates from than Hepaticopsida; when archesporium, it gets differ-
the innermost layer young, the archesporium entiated into spore mother
of amphithecium. It overarches the columella; cells and pseudoelaters.
divides to form 1 to its position is in between The pseudoelaters are ster-
4-layered sporogenous several-layered thick cap- ile, more slender, elliptical
tissue, which is one cell sule wall and the centrally cells with smaller nuclei.
in thickness in A. erectus, located columella.
two-layered in A. pear-
sonii, and 2 to 4 cells in
thickness in A. hallii. The
entire endothecium forms
the central axis in the
form of columella.
7. Sphagnum Amphithecial; arches- In position, it is superficial, Fate of the archesporium
porium originates from as in Anthoceros; when is that entire sporogenous
inner layer of amphith- young, it is dome-shaped tissue develops into spores.
ecium, which divides and and present in the upper part Inside the capsule, this tis-
forms a 4-layered thick of the capsule, overarching sue thus forms a coherent
sporogenous tissue. the tip of the centrally-locat- tract of fertile cells.
ed massive columella within
the wall of the capsule.
8. Funaria Endothecial; archespo- Position of archesporium is Fate of archesporium is that
rium originates from superficial, which surrounds all sporogenous cells are fer-
the outermost layer of the centrally-located colu- tile and form spores; similar
endothecium; a two-cells mella like a barrel. to Sphagnum, the sporog-
thick sporogenous tissue enous tissue forms a coherent
is resulted. tract of fertile cells.
1. Amphithecium The amphithecium either gives rise to (i) capsule wall, as in Hepaticopsida,
Bryopsida and a few species of Notothylas and also Andreaea, or it gives rise to (ii) archesporium and
capsule wall, as in majority of Anthocerotopsida and Sphagnum.
2. Endothecium The endothecium gives rise either to (i) archesporium, as in Hepaticopsida and a
few species of Notothylas, or to (ii) columella, as in most species of Anthocerotopsida and Sphagnidae,
or to (iii) archesporium and columella, as in some Bryopsida and Andreaea.
A NOTE ON THE ORIGIN OF ELATERS 19.5

As mentioned earlier also, elaters are long, thin, sterile cells in the capsule of the sporophyte of a
liverwort. They have spiral thickenings of the cell wall, and have the ability to change their position
with changes in humidity. Elaters help in the dispersal of spores from the capsule.
268 � Bryophyta
Regarding the origin of elaters, some bryologists believe that in Hepaticopsida, certain cells in the
young sporogonium become sporocytes or spore mother cells while others form elaters, and both of
them arise independently from the undifferentiated sporogenous tissue. Other bryologists, however,
opined differently and believe that in some Hepaticopsida (e.g. Pellia, Plagiochasma, etc.) and
Anthocerotopsida (e.g. Notothylas), each undifferentiated cell divides into two daughter cells, of which
one gives rise to one or more spore mother cells and the other develops into one or more elaters. This
later view is supported by Goebel (1927), who also advocated that such a “spore-elater division” of
fertile and sterile cells is true in all elater-containing Hepaticopsida. This is not seen in members where
the sterile cells or elaters are not found, as in Riccia and Sphaerocarpos.
In genera like Targionia and Reboulia, the spore mother cells and elaters may be the members
of the same cell generation or there may exist a difference of 3 to 5 or more generations between
the two. Parihar (1987) mentioned that in Marchantia polymorpha, “the elaters are differentiated five
generations before spore mother cells”. In some other genera, however, elaters are differentiated 3 to 5
generations later than that of the generation of spore mother cells, e.g. Stephensoniella.
1. The first cell genera on of the sporogenous ssue in bryophytes is known as _____.
2. How can you define the term archesporium?
3. What is archesporium? Describe its origin and fate in bryophytes giving suitable examples.
Your descrip on should not exceed 500 words
4. Make a list of at least five products of archesporium.
5. What are elaters? How do they differ from elaterophore?
6. Name a bryophy c genus in which the elaterophore is present at the base of the capsule.
7. In which bryophy c genus is the elaterophore present at the top of the capsule?
8. Two fundamental embryonic layers of the capsule in bryophytes are _____ and _____.
9. Make a table of comparison giving details of origin, posi on and fate of archesporium in
Riccia, Marchan a, Anthoceros and Funaria.
10. The archesporium is _____ in origin in Marchan a.
11. In Anthoceros, the archesporium is _____ in origin.
12. Write a botanical note on the origin of elaters in bryophytes in about 100 words.
20
Apogamy and
Apospory in
Bryophytes
APOGAMY AND APOSPORY:
TWO ALTERNATIVE PATHWAYS OF LIFE CYCLE 20.1
As mentioned earlier also under Article 15.4, apogamy is the phenomenon of the development of
a sporophyte directly from a cell of gametophyte without fusion of gametes so that the resulting
sporophyte has the same chromosome number as the parent gametophyte. On the other hand, apospory
is the phenomenon of the development of a gametophyte directly from a sporophyte cell without
meiosis and the formation of spores. The resulting gametophyte has the same chromosome number as
the parent sporophyte. Some major aspects of the two phenomena (apogamy and apospory), we shall
discuss here in this chapter
All bryophytes exhibit a well-defined heteromorphic alternation of generations, in which the
gametophytic generation is predominant and present in the form of a well-defined thalloid or foliose
gametophore, while the sporophytic generation is dependent on gametophyte and present on it in the
form of sporophyte, usually made up of foot, seta and capsule.
The phenomena of apogamy and apospory, defined above in the first paragraph, are, therefore, two
alternative pathways of life seen in some bryophytes. In nature, however, both these phenomena are
rare, hence their significance in the usual life-cycle process is meagre.
WHY ARE THE PHENOMENA OF APOGAMY AND

APOSPORY OF CONSIDERABLE INTEREST? 20.2
In spite of the rare occurrence of apogamy and apospory in nature, these are of considerable interest for
bryologists because of the following reasons:
1. We can observe and study differentiational processes at the cellular level from the very
beginning because of the studies of apogamy and apospory.
2. We can obtain, preserve and study tissues with haploid and diploid compliments and can use
them in several other experimental studies.
270 � Bryophyta
3. We can study more details of the normal phenomenon of alternation of generations due to the
studies of apogamy and apospory.
4. Callus (a tissue consisting of large, thin-walled, parenchymatous cells developing as a result
of injury, as in wound healing) is usually formed at the beginning of apogamy and apospory.
By modifying the cultural conditions, controlled differentiation of callus into gametophytes or
sporophytes can be achieved and studied.
5. While studying phenomena of apogamy and/or apospory, the role of external factors regulating
or effecting differentiation can be studied more specifically.
6. By maintaining homogenous cell clumps, formed during experimentations of apogamy
and apospory, we can find and develop suitable materials for the production of secondary
metabolites.
7. Biosynthetic problems relating to several compounds can be studied and solved while studying
apogamy and apospory phenomena in bryophytes.
APOGAMY 20.3
20.3.1 Apogamy Occurrence During Diploid and Haploid Phases
Few details of the occurrence of apogamy in diplophase and haplophase are listed below:
1. Springer (1935) was the first to report occurrence of apogamy in sporophytes of mosses. He
reported apogamous sporophytes on the leaf tips of naturally occurring diploid gametophytes
of Phascum cuspidatum. This is the only reported example of apogamy in vivo (in vivo means
biological processes occurring within the living organism or cell). Two alternative paths of
differentiation were shown by the swellings on the leaf tips of P. cuspidatum. When plenty
of moisture was available, protonema was produced in this moss. But in dry conditions or
if increased amount of salt is added in the nutritive medium, these swellings of the leaf tips
developed into apogamous sporogonia.
2. Bauer (1956) reported in Georgia pellucida “that differentiation of apogamous sporophytes is
to a great extent favoured by relative dryness of the nutritive medium”.
3. Bauer (1957) observed in Physcomitrium pyreforme that, instead of protonema, young
sporophytes regenerated to form new sporophytes in this species.
4. In yet another study, Bauer(1959) obtained apogamous sporophytes from the diploid
protonema, regenerated from the intergeneric hybrid sporophyte of Physcomitrium pyreforme
and Funaria hygrometrica.
Apogamy in haplophase has been reported in many mosses including Desmatodon randii by
Lazarenko et al. (1961), Funaria hygrometrica by Chopra and Rashid (1967), Tetraphis pellucida by
Hughes (1969) and Physcomitrium pyreforme by Menon (1974). Lazarenko (1965) reported coexistence
of apogamous sporophytes with normal sporophytes on the haploid plants of Desmatodon randii.
Apogamy in diplophase has been reported in many mosses including Phascum cuspidatum by
Springer (1935), Physcomitrium pyreforme by Bauer (1957, 1959), Desmatodon ucrainicus and D.
randii (Fig. 20.1 A, B) by Lazarenko (1960), Grimmia pulvinata by Hughes (1969) and Funaria
hygrometrica by Kumra and Chopra (1980).
Apogamy and Apospory in Bryophytes � 271
Fig. 20.1 Apogamous sporogonia in diplophase of Desmatodon ucrainicus (A) and

D. randii (B) (A er Lazarenko, 1960)
20.3.2 Produc on of Viable Spores by Apogamous Sporophytes

Apogamous sporophytes of only a few mosses produce the viable spores in haplophase, and one such
case was reported by Lal (1961) in Physcomitrium coorgense. However, many reports are available of
apogamous sporophytes producing viable spores in diplophase in mosses, and some such investigated
species include Phascum cuspidatum by Springer (1935), Physcomitrium pyreforme by Bauer (1959),
Desmatodon randii and D. ucrainicus by Lazarenko (1960), Brachythecium campestre by Lazarenko
(1961) and Pottia intermedia and Splachnum ovatum by Lazarenko (1965).
20.3.3 Produc on of Apogamous Sporophytes from Callus

Several workers, while working on apogamy in bryophytes, have worked on differentiation of
apogamous sporophytes from the callus in different genera including Physcomitrium pyreforme (Bauer
1957, 1961), P. coorgense (Lal, 1961, 1963) and Funaria hygrometrica (Kumra, 1981).
Bauer (1957, 1961) observed that “besides producing protonema, the sporophytes of Physcomitrium
pyreforme, Funaria hygrometrica, and their hybrid sporophytes form calli which later on differentiate
into daughter sporophytes”.
Lal (1961, 1963) observed that the callus obtained from the haploid gametophytic axis and archegonia
of Physcomitrium coorgense also differentiate into apogamous sporophytes. In similar fashion, Kumra
272 � Bryophyta
(1981) observed that the callus obtained from the haploid protonema of Funaria hygrometrica produced
apogamous sporophytes or gametophytes in different environmental conditions.
20.3.4 Factors Responsible for Differen a on of Apogamous Sporophytes

Some major factors which control differentiation of apogamous sporophytes include light, hydration,
sugars, growth hormones, inorganic nutrients, and several endogenous factors (e.g. sporogon factor,
age of tissue, and genetic constitution).
1. Light Diffuse light promotes production of sporophytes and gametophytes from the gametophytic
callus of Physcomitrium coorgense while in darkness, only apogamous sporophytes get differentiated in
this species. Lal (1963) opined that light plays a positive and definite role in establishing the behaviour
of the apical cell. Hughes (1969) observed that in daylight, the frequency of apogamy in the diploid
protonema of Phascum cuspidatum is very low, but it is exceptionally increased in yellow filtered
fluorescent light.
2. Hydra on Springer (1935), the first bryologist to work on apogamy, observed and suggested
that reduced hydration of the medium favours apogamy in Phascum cuspidatum. Observations of
Springer were later on confirmed in some other bryophytes, such as Georgia pellucida (Bauer, 1956),
Desmatodon ucrainicus (Lazarenko, 1960) and Splachnum ovatum (Lazarenko, 1961). Chopra and
Rashid (1967) reported that drying of medium induces apogamy in Funaria hygrometrica.
3. Sugars It has been observed in some mosses (e.g. Physcomitrium coorgense) that sucrose, and
also glucose to some extent, influences the induction of apogamous sporophytes. Rashid and Chopra
(1969) studied and suggested that in Funaria hygrometrica, an enhancement in sucrose concentration
in the medium promotes initiation of apogamous sporophytes from the axis. Kumra and Chopra (1980)
also reported in F. hygrometrica that a 2% addition of sucrose in the medium favours induction of
protonema as well as apogamous sporophytes.
4. Growth Hormones Several such studies have been made on the effect of growth hormones on
apogamy in bryophytes, of which two are on Georgia pellucida and Funaria hygrometrica.
In Georgia pellucida, Bauer (1956) observed that the number of sporogonia per unit area of
protonema increases by adding low concentrations of indole-acetic-acid.
In Funaria hygrometrica, Rashid and Chopra (1969) observed that the “capacity of gametophytes
to produce sporophytes is influenced to varying degrees by growth substances”. If low concentrations
of GA3, IAA, kinetin and a mixture of kinetin and IAA is applied, it “appreciably increases the number
of sporophytes per culture”. If higher concentrations of kinetin is applied, it proves “inhibitory for
gametophyte as well as sporophyte differentiation, but GA3 and IAA inhibit sporophyte formation”
without showing any effect on the growth of gametophyte.
In Physcomitrium pyriforme, Menon (1974) observed that abscisic acid is inhibitory for growth of
protonema and also for the differentiation of apogamous sporophytes”.
5. Inorganic Nutrients Varied types of influences of inorganic salts have been observed on the
induction of apogamous sporophytes in bryophytes. A few such examples are listed below:
Apogamy and Apospory in Bryophytes � 273
(a) Bauer (1959) observed that nitrogen promotes apogamy in Splachnum luteum.
(b) Lal (1964) observed that higher concentrations of salts inhibit the formation of apogamous
sporophytes in Physcomitrium coorgense.
(c) Menon (1974) observed no change in the incidence of apogamy in Physcomitrium pyriforme if
there is a “change in nitrogen source or doubling of the nitrate and chloride concentration”.
6. Endogenous Factors
(a) Sporogon Factor Bauer (1959) was the first to suggest the presence of a factor called sporogon
factor, which emanates from the diploid sporophyte and is translocated into the aposporous protonema.
This factor multiplies in the tissue and is inherited during vegetative propagation. The sporogon factor
appears to be of hormonal nature and may be a mixture of hormones. Application of one such factor
(bryokinin) enhances apogamy in Splachnum ovatum (Bauer, 1966).
(b) Age of Tissue Bauer (1963) suggested that apogamy seems to depend on age of the tissue. He
observed that when sporogonia attain ageing, they regenerate only protonema, irrespective of the
zone.
(c) Gene c Nature It has been observed that chromosome number appears to play an important role
in apogamy. But it also appears that polyploidization is an important factor is apogamy.
20.3.5 Differen a on of Gametophore and Sporophyte

In mosses, usually an apical cell with three cutting faces is established prior to the differentiation of
gametophores, while differentiation of sporophytes is preceded by establishment of an apical cell with
two cutting faces. Apical cell, if present in the protonemal phase, usually contains only one cutting face.
Parihar (1972) and others believe that an apical cell with three cutting faces usually passes through a
stage bearing only two cutting faces. If conditions suitable for apogamy are existing, the apical cell
with two cutting faces gets stabilised and thus develops the sporophyte. On the other hand, if conditions
for its further development into an apical cell with three cutting faces are available, this results into the
differentiation of gametophores.
20.3.6 Role of Calyptra in the Development of Sporophyte

Calyptra is a hood of tissue produced from the wall of the archegonium, especially in mosses. Calyptra
protects the young capsule or sporophyte till the latter is nearly mature. It is usually a membranous,
cap-like body formed after fertilization. Studies of Hughes (1969) and others suggest that “gametophyte
exerts a controlling influence on the developing sporophyte mainly through calyptra”. Rashid and
Chopra (1969) have also proved the significance of the role of calyptra in their studies of apogamous
sporophytes.
APOSPORY 20.4
20.4.1 Apospory and Who Discovered it in Bryophytes?
Apospory (production of diploid gametophyte from vegetative cells of sporophyte without the production
of spores) was discovered by Pringsheim (1876) in Bryum caespitosum, Hypnum cupressiforme and
274 � Bryophyta
H. serpens and Stahl (1876) in Ceratodon purpureus. Wettstein (1925) developed the technique of
seta regeneration for obtaining diploid gametophytes for genetic studies in Funaria hygrometrica,
Physcomitrium pyriforme and some other bryophytic genera.
20.4.2 Apospory in Mosses

Besides the studies of apospory in mosses mentioned above, P Kachroo (1954) studied apospory in
Physcomitrium pyriforme, Narayanaswami and Lal (1957) studied this phenomenon of the regeneration
from the capsule wall in Physcomitrium cyathicarpum, and Kumra and Chopra (1980) studied the
same aspect of seta regeneration in Funaria hygrometrica. It has been observed that the protonema
regenerated from the seta of F. hygrometrica possesses aposporous gametophytes. Kumra and Chopra
(1980) observed that these diploid gametophytes grow very slowly and are much smaller than that of
haploid gametophytes.
20.4.3 Apospory in Liverworts and Hornworts

Many reports of apospory in liverworts and hornworts are, however, not available, but some of these
reports are listed below:
1. Burgeff (1943) studied apospory in Marchantia polymorpha. He developed diploid thalli
aposporously in this liverwort.
2. Matzke and Raudzens (1968) studied apospory in yet another liverwort, Blasia pusilla and
observed almost the same details like that of Marchantia polymorpha.
3. Matzke and Raudzens (1969) reported apospory in two more liverworts (Pellia epiphylla and
Pallavicinia lyellii).
4. Simone (1966) reported apospory in some Jungermanniales, such as Jungermannia lanceolata,
Porella pinnata, Rudula complanata and Lophocolea heterophylla.
5. Aposporous, diploid gametophytes have been obtained from the setae of Athalamia pusilla, a
member of Marchantiales (Mehra and Pental, 1976).
6. Borenhagen (1926) observed aposporous growth in Anthoceros, a hornwort.
1. Write an essay on apogamy and apospory in bryophytes in about 500 words.

2. In bryophytes, two alterna ve pathways of life cycle are _____ and _____.
3. Define the terms ‘apogamy’ and ‘apospory’ in about 50 words.
4. Are the phenomena of apogamy and apospory rare in nature?
5. Apospory and apogamy are of considerable interest for bryologists. Why?
6. What is callus?
7. Give an account of apogamy in bryophytes in about 500 words only.
8. Describe some major factors responsible for differen a on of apogamous sporophytes.
9. What is calyptra? Describe its role in the development of sporophyte in about 50 words
only.
10. Who discovered apospory in bryophytes?
11. Give an account of apospory in bryophytes in about 200 words.
21
Physiology of
Bryophytes
Physiology of an entire, well-established group of plant kingdom and limit of a few pages in a textbook
of this nature, as it is, is not possible, even to summarise. A very elementary account of some of the
selected physiological processes is given here, just to give a very preliminary idea of the physiology of
bryophytes to the readers.
WATER RELATIONS 21.1

In the field of water relations, some attention has been given to absorption and conduction of water
and solutes in the gametophyte and sporophyte of bryophytes. Some work has also been done on the
aspect of remarkable resistance to drought shown by the spores and vegetative cells of the members
of this group. With reference to water relations, bryophytes are very specific because they have very
limited power of withdrawing water from the substrate, they remain attached with. Most of the water
they require for their physiological processes is derived (i) either from the water falling on them, or (ii)
from the water flowing over them.
21.1.1 Absorp on of Water

Haberlandt (1886) studied rapid movement of dye solutions in the central strands of the axis of
Plagiomnium undulatum and Polytrichum juniperinum. It was shown that there is an upward movement
of water from the base of the axis to the leaves, and it can be compared with the transpiration stream
of higher plants. Bowen (1933) observed that “all bryophytes are capable of absorbing water over the
entire surface of thalli or leafy shoot. On the basis of his studies of absorption and conduction of water,
Buch (1947) concluded that bryophytes have two major physiological groups, viz. endohydric and
ectohydric. A third group of ‘myxohydric’ mosses is also present.
1. Endohydric These are the mosses which have a well-developed conducting strand, e.g. Bryum
capillare. Polytrichum commune and Mnium undulatum have the ability to absorb water by their
rhizoids present at the base and transfer the water from the base up to the actively photosynthesising
leaves at the tip. A transpiration stream is present in such mosses. Majority of the endohydric mosses
276 � Bryophyta
are tuft-forming. They have well-developed basal rhizoidal system. They usually do not grow on rocks
or tree barks, but occur frequently on loose substrata, such as soil or humus.
2. Ectohydric These are the bryophytes which lack well-developed conducting strands. Ectohydric
includes all the leafy liverworts and majority of mosses, such as Cryphaea, Orthotrichum, Rhacomitrium
and Ulota. They have the ability to absorb water and dissolved substances through any part of their
external surface (thallus or shoot). They lack regular internal movement of water within their body
parts. Absorption of dew is also very important as a source of water to enable photosynthesis in these
bryophytes.
In ectohydric bryophytes, the leaves often revive within a few seconds while in endohydric
bryophytes, the air-dry leaves become turgid very slowly.
3. Myxohydric In this group of bryophytes, features of both endohydric and ectohydric bryophytes
are present, e.g. Funaria hygrometrica. Such mosses show both external and internal conduction of
water. They occur mainly on moist, porous and nutrient-rich substrata.
21.1.2 Conduc on of Water

For details, refer Chapter 28 (Conduction in Bryophytes).
GROWTH FORMS IN BRYOPHYTES 21.2

Bryophytes have special structural features, due to which their ability to hold capillary water is increased.
They possess specialised growth forms. Some have aggregated shoots and are thus gregarious while
others are solitary and have separated shoots, and many are hanging bryophytes in which the secondary
branches are long and pendulous. A growth-form classification for tropical forest bryophytes has been
proposed by P W Richards (1983). An outline of the same is given here as under:
Table 21.1 Growth-form classifica on for tropical forest bryophytes, as proposed by Richards (1983)
(A) Social Forms Aggregated leafy shoots or thallus branches.

(i) Cushions, in which shoots are mainly erect and radiating to aggregated, cushion-like dome-shaped structures.
(a) Large cushions, e.g. Leucobryum
(b) Small cushions, e.g. Octoblepharum
(ii) Turfs, in which shoots are upright or somewhat parallel.
(a) Tall turfs with mostly erect branches, e.g. Leucoloma.
(b) Tall turfs with divergent or creeping branches, e.g. Sphagnum, Macromitrium
(c) Short turfs less than 2 cm in height, e.g. Diphyscium
(d) Open turfs, e.g. Drepanophyllum
(iii) Mats, in which stems usually creeping over the substratum, forming closely interwoven mats.
(a) Rough mats, e.g. Sematophyllum
(b) Smooth mats, e.g. Radula, Frullania
(c) Thread-like mats, e.g. Lejeunea
(d) Thallose mats, e.g. Dumortiera
Physiology of Bryophytes � 277
(B) Solitary Forms Leafy shoots or thallus branches not aggregated

(i) Protonemal bryophytes, e.g. Ephemeropsis
(ii) Unbranched dendroid forms, e.g. Pogonatum, Dawsonia
(iii) Branched dendroid forms, e.g. Protothanium
(iv) Feather forms, e.g. Bryopteris
(v) Bracket mosses, e.g. Spiridens
(vi) Hanging bryophytes, e.g. Plagiochila, Frullania
WATER HOLDING CAPACITY OF BRYOPHYTES 21.3

In Sphagnum and some oher mosses, the water-holding capacity perhaps depends on the morphological
construction of the shoots. Many bryophytes grow very slowly, and this important factor affects
competetion between them and other surrounding plants, specially higher plants. Temperature and
moisture are also important factors in determining the growth rate and water-holding capacity of
bryophytes.
PHOTOSYNTHESIS 21.4
Haberlandt (1886) was the first to establish that in mosses “all cells of protonema contain enough
chloroplast to be able to assimilate CO2 in the manner of thalli of liverworts and leaves of the green
higher plants”. It has also been observed and suggested that moss leaves and thalli of Marchantiales and
a few other liverworts are better adapted for photosynthesis than many leaves of higher plants. There
appears no major difference in the mode of photosynthesis in bryophytes and higher plants. In most
of the bryophytes, formation of the starch can be observed easily. Instead of starch, some species of
Frullania and Andreaea, however, produce other types of carbohydrates. Starch grains of bryophytes
may contain a large amount of other carbohydrates, including glycogen. Cell walls of many bryophytes
contain hemicellulose and pectin.
Bryophytes require optimum light and temperature for photosynthesis, as required by higher plants.
Coloured substances, found in the cell walls of some liverworts and other bryophytes, are mainly due
to strong light combined with high temperature.
Some major factors, which affect photosynthesis in bryophytes, are availability of water,
temperature and light intensity.
1. Water Content In Hypnum triquetrum, photosynthesis activity increases with increase of water
content up to as much as 300%. When water is supplied to this moss, photosynthesis begins within
5–10 minutes, attains a suitable higher rate within 25 minutes to utilise CO2 produced by respiration,
and within 30-40 minutes it reaches an equilibrium. In Rhacomitrium, very high rate of photosynthesis
occurs in suitable light intensity, when water content is 200–300% of the dry weight and temperature
in the surrounding environment is 12–15°C.
2. Temperature In Rhacomitrium, a moss, the balance between respiration and photosynthesis

is markedly effected by temperature. Photosynthesis rate declines appreciably after a temperature
278 � Bryophyta
of 13–15°C. In Bryum sandberghii, yet another moss, the optimum temperature for photosynthesis
is between 24–30°C but its gametophytes have the capacity to respire and photosynthesise even at
–5°C, as observed by Rastorfer and Higinbothom (1968). According to Atanasiu (1971), the lowest
temperature for photosynthesis in Brachythecium geheebii and Camtothecium philippeanum is –9°C,
and for Isothecium viviparum is –8°C, and all these three are mosses.
3. Light Intensity In epiphytic mosses, the intensity of light plays a definite role on the rate of
photosynthesis. It has been proved that the upper limit of vertical distribution of these mosses on
trees of forests is restricted primarily by water but their lower limit is restricted by light intensity.
The chlorophyll content and photosynthetic efficiency of epiphytic mosses resembles those of higher
green plants. It has been shown by Hahn and Miller (1966) in Polytrichum commune that chloroplast
replication in this moss takes place “in continuous white light and red light of 15 minutes/6 hours. In
continuous darkness and in far-red light of 15 minutes/6 hours, the size of chloroplasts increased” but
there is no effect on their number.
NITROGEN METABOLISM 21.5

Schuler et al. (1955), Diller (1966) and Montagne et al. (1969) are some of the workers who have worked
on nitrogen metabolism in bryophytes. Montagne et al. have studied the enzymatic transamination by
Sphaerocarpos texanus cultivated in vitro. Their studies show similarity between the transaminase
system of this liverwort (S. texanus) and that of angiosperms.
RESPIRATION 21.6
Although much work has not been done on respiration in bryophytes, two major workers who have
reviewed this aspect of bryophytes are A J M Garjeanne (1932) and W Stiles (1960). Instead of respiration,
most researchers have focused on the capacity of the bryophytes to survive severe desiccation, and also
on the relationship between respiration and water content. However, it has been established that there
is no difference between the mode of respiration in bryophytes and that of higher plants. The rate and
intensity of respiration in bryophytes is also affected by all the factors in almost all the same ways as
that of higher plants. These factors include light, temperature, amount of available oxygen, the amount
of carbohydrates in the cells, amount of CO2 in the surrounding atmosphere, concentration of salts in
the soil, etc.
It has been observed in Sphagnum and some other mosses that weak solutions of nutritive salts
increase the intensity of respiration. The respiration process diminishes constantly in Sphagnum, if kept
in distilled water. Solutions containing calcium salts first show an increase in rate of respiration but
later on they show negative effect because its pH becomes unfavourable for the process.
Thalli, or leafy parts of several bryophytes, live in close contact with the soil. Air in such surroundings
is relatively rich in CO2 and water, and O2 is in lower amounts. Several microorganisms, like algae,
fungi, bacteria, protozoa, etc. occur freely in such surroundings. In such environment, the respiration
of bryophytes is lowered. However, the process of respiration goes on regularly.
RQ (Respiratory Quotient the ratio of moles CO2 evolved to moles O2 absorbed in respiration)
of many mosses including Polytrichum juniperum, Hypnum triquetrum, etc. have been studied by
Physiology of Bryophytes � 279
Bastit (1891) and Plantefol (1927) and it was suggested that RQ. is that of a carbohydrate substrate
irrespective of whether the plants are turgid or in a state of partial desiccation.
Studies have proved that effect of temperature on the respiration in mosses is almost the same as that
in higher plants. In the germinating spores of Polytrichum juniperum, it has been shown by Paolillo and
Jagels (1969) that respiration is limited by hydration rising to a maximum after 10 minutes. In some
mosses, it has been shown that respiration and photosynthesis are increased in autumn but decreased
in winter and reach their lowest level in February and again increase in the coming spring season
(Atanasiu, 1971).
ENZYMES 21.7
Enzymes are usually large, complex protein molecules which, in very small quantities catalyse and
control the natural chemical reactions of metabolism. In bryophytes, Udar and Chandra (1960) detected
in male and female plants of Riccia discolor several enzymes, including aldolase, amylase, butyrase,
catalase, deaminase, invertase, lipase, maltase, phosphatase, phosphorylase, ribonuclease and urease.
In yet another study in 1960, these two Indian bryologists reported several enzymes in four other
liverworts and opined that “metabolic processes in hepatics may closely correspond to those obtained
in green tissues”.
Maravolo et al. (1967) studied biochemical changes during sexual development in Marchantia
polymorpha and reported phosphatases, esterases and peroxidases from the extracts of different parts
of its thalli, including antheridiophores and archegoniophores.
Rao et al. (1969) studied the behaviour of oxidising enzymes of the thalli of Riccia plana by
subjecting the plants to different periods of light and darkness. It was noticed that the activity of
ascorbic acid oxidase is higher in plants grown in darkness. On the basis of their studies, these workers
suggested that “light has an inhibitory effect on the activity of the enzymes.”
Taylor et al. (1970) separated many forms of formic dehydrogenase, glutamic dehydrogenase, lactic
dehydrogenese and malic dehydrogenase from 20 species of bryophytes.
ORGANIC ACIDS 21.8

Allsopp (1951) was the first to make a chromatographic survey of eight liverworts but he could not
detect any acids in them. Ten years later, VSR Das in 1961 reported large quantities of aconitic acid
in Riccia. Das and Rao (1966) detected mannuronic acid in the cell wall of Riccardia. Detection of
organic acids in bryophytes still needs much work to be done.
MOVEMENTS 21.9
Responses to stimuli of the bryophytes awaits a lot of work still to be done. Garjeanne (1932) published
a brief account of such studies in bryophytes in Fr. Verdoorn’s Manual of Bryology. A detailed account
of tropisms and other movement, specially geotropic responses of Marchantia, has been given by Miller
and Voth (1962), published in Volume 65 of Bryologist.
280 � Bryophyta
Spermatozoids of bryophytes show movement, and this is a specific property of many of them.
Chemotropism (growth of plant organ in response to a chemical stimulus) and hydrotropism (growth
in response to the stimulus of moisture) is shown by spermatozoids of bryophytes. Researches have
shown that chloroplasts of bryophytes exhibit phototactic responses. It was Rawitscher (1932) who
reviewed the researches regarding the effect of gravity, light and other factors on the thallus and
tissues of Marchantia. Kohlenbach (1957) performed several experiments on geotropic response of the
gemmae in Marchantia.
1. Give a brief account of some major events of physiology of bryophytes in about 500 words.
2. How can you differen ate between endohydric, ectohydric and myxohydric bryophytes?
3. The bryophytes which lack well-developed conduc ng strands are known as _____.
4. Give an example of a myxohydric moss.
5. Enumerate some major types of growth forms in bryophytes.
6. Give an example for protonemal bryophyte.
7. What are hanging bryophytes?
8. Give an example of a hanging bryophyte.
9. Describe some developments in the field of photosynthesis in bryophytes in about 250
words.
10. Write a brief scien fic note on respira on in bryophytes.
11. What are enzymes? Write a brief note on enzymes of bryophytes.
22
Chemical
Constituents of
Bryophytes
Bryologists from different parts of the globe have reported several organic compounds from bryophytes
in the recent past. These include antibiotics, terpenoids, lignins, flavonoids, lipids, sterols and growth
substances. In several cases, these have also been utilised in taxonomic categorisation of bryophytes.
ANTIBIOTICS 22.1
Antitumor elements have been reported from the extracts of Marchantia polymorpha, M. stellata and
Polytrichum commune by Hartwell (1971). Antimicrobial activity is shown by several bryophytes.
Dumortiera hirsuta and Conocephalum conicum are active against the fungus Candida albicans while
Sphagnum strictum inhibits the growth of several bacteria including Pseudomonas aeruginosa and
Staphylococcus aureus. Sphagnum inhibits the growth of bacterium Sarcinia lutea (Ramaut, 1959).
Mosses, such as Anomodon rostratus and Orthotrichum rupestre inhibit the growth of several species
of bacteria like Micrococcus and Streptococcus. A pronounced effect over the activity against several
bacteria is shown by several mosses including Sphagnum, Polytrichum and Atrichum (McCleary and
Walkington, 1966). Gupta and Singh (1971) reported antibacterial activity of petroleum ether extracts
of Barbula and Timmiella against 33 species belonging to both Gram +ve and Gram –ve bacteria.
Antibacterial activities of 52 species of bryophytes have been reported by Banerjee and Sen (1979).
According to these workers, however, none of the 52 investigated species of bryophytes possessed
antifungal property. They also opined that antibiotic activity of bryophytes varies from species to
species and it also depends on the age of the plant, season of collection and ecological niche. Some
species of Marchantia and Plagiochasma also have antibiotic activity and cause inhibition of several
bacteria such as Bacillus subtilis, Vibrio cholerae, Sarcina lutea and Staphylococcus aureus. Chopra and
Kumra (1988) have mentioned that the “taxa Campylopus laetus, Asterella sanguinea, Plagiochasma
appendiculatum and Reboulia hemispherica were moderately active against the Gram –ve bacteria,
Salmonella typhi and Vibrio cholerae”. Several bryophytes have antifungal property, e.g. Sphagnum
portoricense and Dumortiera hirsuta (McCleary et al., 1960). Wolters (1964) also studied antifungal
activity of two Jungermanniales and 16 mosses.
282 � Bryophyta
GROWTH SUBSTANCES 22.2

Five major groups of growth substances include auxins, gibberellins, cytokinins, abscisic acid and
ethylene, of which the first three are generally growth promoters while the last two are generally
growth inhibitors.
1. Auxins Chromatographic and some other studies of Narayanaswami and LaRue (1955) suggest
the production of indole-acetic-acid, an auxin, in the apical parts of the thallus of some liverworts,
e.g. Lunularia cruciata. Schneider et al. (1967) and Maravolo (1981) reported the occurrence of IAA
in Marchantia polymorpha and some other bryophytes. Presence of IAA in the gametophytes of
Physcomitrella patens has also been reported by Ashton et al. (1985).
2. Gibberellins Gibberellin-like substances have been reported in some bryophytes like Polytrichum
commune (Muromtsev et al., 1969) and Marchantia polymorpha (Melstrom et al. 1974).
3. Cytokinins Bryokinin, an endogenous cytokinin has been isolated from the callus cells of the
hybrid sporophyte (Funaria hygrometrica X Physcomitrium pyriforme) by Bauer (1966). This cytokinin
is physiologically active at several stages of the development in mosses. It promotes activities like
bud formation, archegonium differentiation and formation of apogamous sporogonium in Splachnum
ovatum.
4. Abscisic Acid and Lunularic Acid Most of the liverworts lack abscisic acid. It has been
reported that the same growth regulating functions, as of abscisic acid, are fulfilled by lunularic acid in
most of the liverworts.
5. Ethylene DeGreef et al. (1981) and Thomas et al. (1983) reported production of ethylene in
Marchantia. It has also been reported in Funaria hygrometrica by Rohwer and Bopp(1985). In mosses,
ethylene works as a senescence hormone.
LIPIDS 22.3
Lipids are a group of chemical compounds which contain glycerol and fatty acids. They are insoluble
in water and soluble in organic solvents. Triglycerides, wax, steryl esters, phospholipids and glycolipids
are some lipids. Of these, considerable amounts of triglycerides, wax, esters and steryls have been
reported in green moss protonema, shoots and spores by several workers including Lilgenberg and
Karunen (1978), and Karunen and Mikola (1980).
ALKANES 22.4
Alkanes are saturated hydrocarbons, i.e. compounds containing carbon and hydrogen. Bryophytes
contain a wide range of alkanes. A detailed list of major alkanes found within several members of
Hepaticopsida and mosses is given by Chopra and Kumra (1988).
Chemical Cons tuents of Bryophytes � 283
FATTY ACIDS 22.5

A fatty acid is an organic acid with the general formula CnH2nO2. Fatty acids have been reported
in several members of Hepaticae including Pellia (Matsuo et al., 1971), Marchantia polymorpha
(Gellerman et al., 1972), Asterella (Caldicott and Eglinton, 1976) and Conocephalum (Matsuo et
al., 1980). Huneck (1983) isolated fatty acids from about 30 genera of mosses including Distichum,
Fontinalis, Hedwigia, Hypnum, Polytrichum, Sphagnum and Tortula. Lipid content of three species of
Sphagnum, at different ages of 1–5 years, have been studied by Karunen et al. (1979).
TERPENOIDS 22.6
The compounds made up of two or more isoprene molecules, i.e. CH2 = C(CH3) – CH = CH2 are called
terpenoids. On the basis of such C5 units present in them, these are classified into various categories
like monoterpenoids containing C5 units (reported only in a few bryophytes, e.g. Radula complanata
and Conocephalum conicum), sesquiterpenoids containing C15 units (e.g. Bazzania trilobata),
diterpenoids containing C20 units (e.g. Jungermannia infusca), triterpenoids and sterols containing
C30 units (e.g. Thamnium alopecurum), etc. As many as 14 steroids have been isolated from bryophytes
as mentioned by Chopra and Kumra (1988), mostly from mosses (e.g. Polytrichum and Sphagnum;
Marsili et al., 1972), hornworts and liverworts (e.g. Anthoceros and Marchantia; Asakawa et al., 1981).
Chopra and Kumra (1988) have given a list of ninety-nine bryophytes containing steroids.
FLAVONOIDS 22.7
Flavonoids are natural phenolic products found in several groups of green plants, including bryophytes.
Flavonoids in the form of anthocyanin-like pigments have been reported in several species of Sphagnum.
Three rare flavonoids have been reported in Bryum cryophilum, a moss, by Benz et al. (1962). Markham
et al. (1969) reported flavonoids from Hymenophyton flabellatum. Flavonoids seem to be more frequent
in Hepaticae than in Musci, and “nearly all investigated species of Marchantiales possess flavonoids”
(Chopra and Kumra, 1988).
Flavones are the predominant type of flavonoids found in bryophytes, e.g. Marchantia polymorpha,
Plagiochila asplenioides, etc. Other flavonoids reported from bryophytes include isoflavones (e.g.
Bryum capillare), flavonols (e.g. Reboulia hemisphaerica), dihydroflavones (e.g. Riccia crystallina),
aurones (e.g. in antheridiophores of Marchantia polymorpha), chalcones (e.g. Plagiochasma rupestre,
P. tenue; Schier, 1974), anthocyanins (e.g. Bryum cryophilum) and sphagnorubins (e.g. Sphagnum
magellanicum).
LIGNINS 22.8
Lignin is a complex aromatic compound which is deposited in the cellulose cell walls of the xylem
and sclerenchyma during the process of secondary thickening. Wood is made mostly of lignin. In
284 � Bryophyta
bryophytes, Siegel (1962) found evidence for the presence of traces of lignin-like materials in peristomial
teeth tissues of Polytrichum and also in the gametophyte axes of giant mosses like Dawsonia and
Dendroligotrichum. Presence of lignin, built mainly of p-hydroxyphenyl units, has been shown by
Bland et al. (1968) in Sphagnum. However, the presence of true lignin in bryophytes is still doubtful
and needs further investigations.
CAROTENOIDS 22.9
Using paper chromatography methods, Douin (1956, 1958) investigated the carotenoids of 20 species
of Marchatiales and Jungermanniales and 40 species of Bryales as well as some species of Andreaeales
and Sphagnales, and concluded that a-carotene, b-carotene and lutein are present in almost all these
taxa. Along with these carotenoids, some others like violaxanthin and zeaxanthin have also been
reported in some species of mosses by Freeland (1957). As many as ten carotenoids have been isolated
from Fontinalis antipyretica by Benz et al. (1968). In spores of Polytrichum commune, Karunen and
Ihantola (1977)) reported the presence of carotenes, violaxanthin, lutein, neoxanthin and zeaxanthin.
Chopra and Kumra (1988) have given a detailed list of carotenoids found in 9 genera of liverworts and
15 genera of mosses.
CARBOHYDRATES 22.10
Several types of carbohydrates have been reported from the bryophytes including glucose, fructose
and sucrose, as in Sphagnum. As many as 40 genera of liverworts and 6 genera of mosses have been
investigated for this purpose. Some major carbohydrates met within some common genera of liverworts
include raffinose (e.g., Diplophyllum), sucrose (Fossombronia), amidon and hexitol (Marchantia),
inuline (Monoclea), amidon and maltose (Pellia), mannuronic acid (Plagiochasma), and glucose
and fructose (e.g. Porella, Reboulia). Amongst the mosses, glucose, maltose and sucrose, along with
small amounts of mannose, melibiose and deoxyribose have been reported in Torula, Rhyncostegium,
Platyhyphidium and Homalothecium (Margaris and Kalaitzakis, 1974).
ORGANIC ACIDS 22.11

Much work has still not been done on organic acids in bryophytes. Das and Rao (1963) have shown
the presence of transaconitic acid and malic acids in Riccia, Plagiochasma and Riccardia. The latter
two genera also contain mannuronic acid. Citric acid, fumaric acid, malic acid and succinic acid have
been reported in Sphagnum by Maass and Craigie (1964). Derivatives of cinnamic acid have been
reported to occur in several bryophytes including Sphagnum magellanicum (Tutschek et al., 1973) and
Anthoceros (Mendez and Sanz-Cabanilles, 1979). Some other organic acids have also been reported
from some bryophytes like cis-aconitic acid (Riccia), glycolic acid (Plagiochila), and shikimic acid
(Pellia conocephalum).
Chemical Cons tuents of Bryophytes � 285
ENZYMES 22.12
Enzymes have been reported to occur in several bryophytes. Amongst the liverworts, Udar and Chandra
(1960) reported amylase, butyrase, catalase, invertase, laccase, lipase, maltase and urease from Asterella
and Plagiochasma. Marchantia contains esterase, urease and phosphatase, according to Jones et al.
(1973) while Lophocolea and Frullania contain urease and oxalic acid oxidase, respectively. Amongst
mosses, enzymes have been reported in over 30 genera, of which major ones include Dawsonia,
Fontinalis, Funaria, Hypnum, Pogonatum, Polytrichum, Sphagnum and Tortula (Chopra and Kumra,
1988).
AMINO ACIDS 22.13

Amino acids within bryophytes have been investigated in only a few members. Amongst the Hepaticae,
they have been investigated in Marchantia (Schneider et al., 1967), Pellia, Plagiochila and Diplophyllum
(Touffet and Villeret, 1958). Some amino acids reported from these genera include tryptophan, allantoin,
alanine, aspartic acid, glycine, glutamic acid, lysine, proline, valine, etc. Amongst the mosses, they have
been investigated in Funaria hygrometrica and some species of Sphagnum (Hartmannn and Geissler,
1973; Black et al., 1955). Some of the amino acids reported from these mosses include allantoine,
allantoic acid, arginine, glycine, histidine, leucine, lysine, proline and tyrosine.
ALLERGENIC AND ANTI TUMOUR ACTIVITIES 22.14

Liverworts like Radula and Frullania cause skin allergies, and the active allergenic principle in some
species of these genera is sesquiterpene lactone frullanolide according to Perold et al. (1972).
Potential allergenic agents have been reported from some members, such as (i) germacranolide from
species of Frullania and Porella, and (ii) eudesmanolides from Diplophyllum.
In the treatment of Sarcoma-37, a type of malignant tumour of connective-tissue origin, ethanolic
extract of Polytrichum juniperum has proved to be effective (Belkin et al., 1953). A compound
diplophyllin has been isolated from Diplophyllum. This is effective against human epidermoid
carcinoma according to Ohta et al. (1977).
1. Write an account of the chemical cons tuents of bryophytes in about 500 words.
2. Name any five bryophy c genera which have an microbial ac vity.
3. Write a note on growth substances in bryophytes in about 200 words
4. Name a member of Marchan ales, from which at least three growth substances have been
reported.
5. Name two bryophytes from which fa y acids have been reported.
6. What are flavonoids? Give an account of flavonoids from bryophytes.
7. “Some bryophytes have allergenic and an -tumour ac vi es”. Comment in about 100 words.
8. Marchan a is the most important genus from the point of view of chemical cons tuents.
Elaborate.
23
Bryophytes as
Indicators of
Environmental
Conditions and
Pollution
Most people, in general, think that bryophytes are of no use to mankind. However, besides their several
economic and ethnic uses (discussed in detail in Chapter 32), bryophytes are of definite ecological uses,
and they are of particular utility as indicators of environmental conditions and pollution. In bryophytes,
pollutants (i) inhibit sexual reproduction, (ii) reduce photosynthesis, and (iii) reduce growth of plants
and may eventually cause their death. Increased pollution has made some species of bryophytes “rare”
and others have even become “extinct”. In general, however, “rough mats, tall turfs, large cushions, and
leafy liverworts are least resistant to pollutants” (Gilbert, 1970). From the point of view of indicators
of pollution, there are two types of bryophytes. The first are extremely sensitive to pollution and
show clear visible symptoms of injury even in the presence of very little quantities of pollutants. Such
bryophytes serve as good bioindicators of the degree of pollution. The second type of bryophytes have
the capacity to absorb and retain pollutants in quantities much higher than those absorbed by members
of other plant groups growing in the same habitat.
INDICATOR SPECIES OF BRYOPHYTES 23.1

Many bryophytes are good indicators of environmental conditions. Terrestrial bryophytes in several
countries, including Finland, are widely used as indicators of specific forest types. Importance of
bryophytes as indicators of several minerals was established in the middle of the 20th century, and it
was R R Brooks (1972) who recommended bryophytes as guides to mineralisation. D C Smith (1976)
worked on indicator species of bryophytes with reference to minerals and proved that “bryophytes could
solve several difficulties that are often associated with stream sediment sampling”. Copper mosses (e.g.
Bryophytes as Indicators of Environmental Condi ons and Pollu on � 287
Scopelophila, Mielichhoferia elongata and M. mielichhoferi), grow almost exclusively in areas high in
copper, particularly in copper sulphate” (Shacklette, 1984).
According to Glime and Keen (1984), presence of Fontinalis and Brachythecium rivulare indicate
the presence of iron oxide in the area. It has been proved by Shiikawa (1962) that Jungermannia
volcanicola, Polytrichum and Sphagnum “play active roles in deposition of iron ore” in any area.
Dierssen (1973) established that presence of Sphagnum is a reliable indicator of acidic conditions in
the surrounding.
Mentioned below are some examples of bryophytes which are indicators of environmental conditions
and pollution:
1. Ceratodon purpureus indicates good drainage and high amounts of nitrogen.
2. Pleurozium schreberi and Pogonatum alpinium are indicators of less nitrogen.
3. Funaria hygrometrica and Pohlia cruda in an area indicate good base saturation.
4. Psilopilum laevigatum is an indicator of poor physical soil condition and poor base
saturation.
5. Absorption of rain and atmospheric water by bryophytes make some of them as pH indicators,
e.g. Polytrichum is a good acid indicator.
6. Presence of Leucobryum in a region is an indicator of acidic soil combined with dry, infertile
and deep humus.
7. J A Janssens (1988) and others have used bryophytes (e.g. Sphagnum) as indicators to identify
past climates. Such studies enhance our information about the bryophytic flora of the past.
EROSION CONTROL BY BRYOPHYTES 23.2

Conard (1935) suggested and proved that “sowing spores and vegetative fragments of bryophytes on
bare areas could help to prevent erosion”. He opined that bryophytes (e.g. Barbula, Bryum and Weissia)
are major “pioneers on new roadbanks, helping to control erosion” prior to the large-sized plants getting
established there. Ando (1957) worked on erosion control by bryophytes and suggested that in Japan,
the bryophytes (e.g. Atrichum, Blasia, Nardia, Pogonatum and Pohlia) “play a role in preventing erosion
of banks” of the rivers, streams and other water reservoirs. Oechel and Sveinbjornsson (1978), on the
other hand, suggested that “when bryophytes such as Sphagnum reach saturation, they can suddenly
release a great load of water at unexpected times. Because of its tremendous water-holding capacity,
Sphagnum along with Calliergon sarmentosum, controls water during spring runoff in the Arctic”.
NITROGEN FIXATION 23.3

Usually, the nitrogen is a limiting nutrient for plant growth, in general, and agriculture in particular.
Because of the presence of nitrogen-fixing blue-green algae (e.g. Anabaena, Nostoc) in many bryophytes
(e.g. Anthoceros), they can contribute significant soil nitrogen, particularly to dry range land soils.
The blue-green algae present in Anthoceros behave symbiotically, take nitrogen from the atmosphere
and convert it to ammonia and amino acids. The extra amount of nitrogen is released to the substrate
where it is usually used by other organisms. The bryophyte crusts provide homes for nitrogen-fixing
organisms, according to Harper and Marble (1988).
288 � Bryophyta
Some bryophytes (e.g. communities of Sphagnum) show very high nitrogen-fixing rates. In this
way, bryophytes function as substrate for nitrogen-fixing organisms, and are thus very important to the
forestry industry. According to Granhall and Hofston (1976), following three types of nitrogen-fixing
associations exist in Sphagnum and other taxa:
1. Epiphytic Cyanobacteria,
2. Intracellular Cyanobacteria, and
3. Nitrogen-fixing bacteria.
Rao and Burns (1990) opined that “nitrogen-fixing Cyanobacteria of bryophyte species also provide
growth enhancement for oilseed rape.”
BRYOMETER OR MOSS BAG 23.4

It is now an established fact that bryophytes have played a major role in maintaining changes in the
atmosphere of the earth. For monitoring these changes, scientists in Japan have developed an instrument
called bryometer. The bryometer, developed by H Taoda of Japan in 1976, is “a bag of mosses that
responds in predictable ways to various levels of air pollution”. Taoda (1976) exposed a variety of
mosses to various levels of SO2 and confirmed that most species of mosses “are injured by 10–40 hours
of exposure at 0.8 ppm SO2, or at 0.4 ppm after 20–80 hours.”
Bryometers are now used throughout the world to monitor changes in the atmosphere. In many
European countries, the bryometer is now used so commonly that it is now named a moss bag.
Monitoring of heavy metals around coal-fired plants in Finland is done by moss bags made of
Hylocomium splendens.
SO2 AND ACID RAIN 23.5

Mosses are not widely used as reliable indicators of environmental conditions. Grimmia pulvinata and
several other bryophytes are used as indicators of SO2 in England and several countries of Europe and
North America. H Taoda (1972) of Japan used several epiphytic species of bryophytes to assess pollution
impact in the city of Tokyo, and “divided the city into five zones, based on pollution intensity”. He
listed four groups of bryophytes “in order of increasing sensitivity to SO2”. In 1980, Taoda used three
bryophytes (Conocephalum supradecompositum, Lunularia cruciata and Marchantia polymorpha) “to
assess the degree of urbanization in Chiba city near Tokyo”.
SO2 fumigation affects mosses and shows reduction in their coverage. It is, however, very difficult
to estimate whether the damage to mosses is directly due to SO2 or if it is the result of the ultimate
formation of sulphuric acid, formed due to acid rain. According to the researches of Raeymaekers
(1987), “acid rain, resulting from SO2 emissions, can actually improve conditions for Pleurozium
schreberi in some jackpine (Pinus banksiana) forests. P. schreberi grew faster and increased in cover
when sprayed with water acidified to pH 4.5”. “However, at pH 3.5, its growth and chlorophyll content
were reduced and capsule production decreased”. It has also been established that “ a pH as low as
3.5 is not uncommon in acid fog. While acid rain may favour some bryophytes, acid fog can be more
damaging.”’
According to Winner et al. (1978), bryophytes can not only serve as warning systems, but can
also “protect the nutrients and roots beneath them. By intercepting sulphate ions, bryophytes prevent
formation of sulphuric acid that contributes to leaching valuable nutrients from soil”.
BRYOPHYTES AS BIOINDICATORS OF
HEAVY METALS IN AIR POLLUTION 23.6
Gilbert (1969) was the first to strongly recommend the “use of cryptogamic epiphytes as biological
pollution indicators”. Since then, the “bryophytes have been used to monitor airborne pollution caused
by emissions from factories”. Scientists in different parts of the world have now strongly correlated the
absence of epiphytic mosses, lichens, and majority of liverworts from urban areas with air pollution.
Rao and Le Blanc (1967), Le Blanc (1969), Rao et al. (1977), Ferguson and Lee (1978), Rao (1982)
and others have now conclusively proved that air pollutants, especially heavy metals, affect growth and
reproduction of bryophytes and lichens.
Bryophytes, particularly mosses and liverworts, suit well as bioindicators and biomonitors because
of their characteristics such as (i) lack of significant cuticle, (ii) lack of significant epidermis, (iii) lack
of well-organised leaves, (iv) “leaves” being only one cell thick, and (v) absence of a well-developed
conduction system. Due to all these characteristics, bryophytes absorb both nutrients and pollutants
directly from the atmosphere.
Scientists have now proved that bryophytes easily “absorb heavy metals without the regulation
characteristic of their nutrient absorption”. This ability of many bryophytes to separate or set apart
“heavy metals while remaining unharmed makes them good biomonitors” or bioindicators. For
example, “Marchantia polymorpha accumulates lead” (Briggs, 1972) and “Calymperes delessertii is a
good monitor for aerial lead and to a less extent, copper” (Low et al., 1985). Nash (1972) has proved
that Pottia truncata, Polytrichum ohioensse, Dicranella heteromella and Bryum argenteum are very
tolerant of high tissue levels of cadmium (610 ppm), copper (2700 ppm), and zinc (55,000 ppm).
Thomas (1983) opined that “Hypnum cupressiforme accumulates three times as much zinc, copper and
cadmium as do lichens or seed plants”.
Satake et al. (1989) investigated that bryophytes have a variety of means by which they can separate
or set apart “substances that are toxic to many higher plants and animals”.
LeBlanc and Rao (1973) proved that in many countries, including Germany and Canada, “bryophytes
have been transplanted from pollution-free areas to areas suspected of pollution damage”.
Studies suggest that heavy metals constitute a very important class of pollutants. The most significant
among these are (i) lead, (ii) cadmium, (iii) zinc, and (iv) mercury. A few examples to prove this are
mentioned below:
1. Lead The most toxic metal known is lead. It tends to accumulate more in ectohydric mosses like
Dicranella varia. These mosses lack cuticle and are, therefore, able to absorb water from their entire
body surface. In Grimmia deniana, lead is ionically bound to the cell wall, and thus prevents its toxic
amount from penetrating the cell wall. Studies of Briggs(1972) in Marchantia polymorpha proved that
lead tolerance can play a major role in natural selection
2. Cadmium Mosses are able to take up airborne cadmium. They can also absorb cadmium
from the substrate. Bryophytes show sensitivity to cadmium at very low concentrations, and even as
290 � Bryophyta
low as 10–8 M. The indications of damage to the bryophytes (e.g. Sphagnum) are noticeable in the
pigmentation and growth rate. Cadmium at 1 ppm can significantly enhance the elongation of germ
tubes in Funaria hygrometrica. At higher concentrations (e.g. 10 ppm), however, it can significantly
reduce spore germination in this moss.
3. Zinc According to Shimwell and Laurie (1972), zinc uptake is related to the water economy
of mosses, e.g. Dicranella varia. In Funaria hygrometrica and Marchantia polymorpha, zinc
concentrations more than 50 ppm result in complete inhibition of spore germination and extension of
germ tube.
4. Mercury Bryophytes collect mercury through rainwater or dust particles. Mosses have the
ability to tolerate high concentrations of mercury, and have, therefore, been widely used as indicators
of atmospheric mercury. Mondano and Smith (1974) recorded 4.34ppm mercury in Dicranella
heteromella in urban regions, but only 0.24 ppm mercury in rural areas. Mercury values in mosses and
some other plants indicate that bryophytes give a more reliable indication of mercury fall-out.
COPPER MOSSES AS INDICATORS OF

COPPER CONCENTRATION 23.7
Some species of mosses serve as indicators of high copper concentration in the substrate, and these
are known as copper mosses, e.g. Dryptodon stratus, Merceya ligulata, Mielichhoferia elongata,
M. macrocarpa and M. mielichhoferi. Some liverworts whose principal substrata are copper ores are
Cephaloziella phyllacantha and Gymnocoba acutiloba. According to Warncke (1968), Marchantia
alpestris in Scandinavia “is generally restricted to copper-rich areas,” and it should be added to the list.
According to Shacklette (1967), many copper mosses grow on substrata low in sulphur, and others on
substrata with relatively high pH.
Copper mosses occur on soil rich in copper but they cannot be used as plant indicators in determining
the presence of copper in the region. It is so because (i) these mosses are very rare, and (ii) it is very
difficult to identify them.
BRYOPHYTES AS BIOINDICATORS OF SOME

OTHER AIR POLLUTANTS 23.8
Besides heavy metals, SO2 and acid rain, discussed already in the earlier part of this chapter, bryophytes
are also useful indicators for other types of air pollution, e.g. hydrogen fluoride and ozone. A bryophyte
sensitive to hydrogen fluoride is Orthotrichum obtusifolium. On the other hand, some bryophytes,
tolerant of fluoride fumes, are Rhacomitrium, Polytrichum commune and P. strictum. According to
Gagnon and Karnosky (1992), some species of Sphagnum are especially susceptible to ozone. They
show characteristics such as (i) reduced photosynthesis, (ii) reduced growth, (iii) loss of colour, and
(iv) symptoms of desiccation. Lee at al. (1998) proved that well-hydrated bryophytes are not generally
sensitive to ozone at concentrations likely to occur in the atmosphere.
BRYOPHYTES AS INDICATORS OF UV B RADIATION 23.9

According to Hedenas (1991), Bryum argenteum, a moss, is now being used widely “to monitor the
thickness of the ozone layer over Antarctica. As the ozone layer decreased, increased exposure to
UV-B (280–315 nm or UV-Medium) radiation stimulated production of flavonoids” in B. argenteum.
However, in Sphagnum magellanicum, there were no noteworthy differences in chlorophyll or
carotenoid concentrations following UV-B exposure, as observed by Searles et al. (2002). Wilson et
al. (1998) reported earlier that “in the presence of adequate water, growth of Hylocomium splendens
in Norway was strongly stimulated by UV-B equivalent to 15% reduction in ozone.” A decrease in
concentration of UV-B-absorbing compounds is also shown by Polytrichum commune after the third
year of its growth.
BRYOPHYTES AS INDICATORS OF RADIOACTIVITY 23.10

Bryophytes have the ability to sequester minerals, and then also they remain unharmed. It is this
ability of bryophytes which makes them “good indicators of accumulated radioactivity” according to
researchers like Whitehead and Brooks (1969), Beckett et al. (1982) and Summerling (1984). Due to
the cation exchange activity of several species of Sphagnum, they “could be used to decontaminate
water containing radioactive materials” (Fischer, et al. 1968). It has been proved by Kulikov et al.
(1976) that “the uptake of radioisotopes by epigean mosses occurs not so much from substrates as
directly from atmospheric fall-out.”
BRYOPHYTES AS INDICATORS IN AQUATIC HABITATS 23.11

Several bryophytes are of specific importance as indicators in aquatic habitats. One of their major
advantage is their ability to integrate pollution over time and keep a record that cannot be achieved
through testing of water chemistry. It is so because their contaminant content is more consistent than
that of the sediments. Bryophytes are used as aquatic bioindicators because (i) they are easy to collect,
(ii) they are easy to transplant, (iii) they can be harvested any time of the year, and (iv) their samples
can be kept for many years for further analysis. Some such suitable bryophytes include species of
Fontinalis, Leptodictyum, Platyhypnidium and Scapania. Scapania undulata can survive at a low pH
of 3.9, and it is a very useful accumulator for zinc, lead and cadmium in nutrient-poor water (Satake
et al, 1989).
In Pohlia ludwigii, a moss, it has been reported by Soma et al. (1988) that “aluminium, manganese,
copper, zinc and lead were in higher concentrations 1–3 cm below growing stem tips than at tips, but
sodium, phosphorus, calcium and iron differed little between the 1 cm tip portion and lower parts.”
This study proves that accumulations differ in different parts of mosses.
Mosses prove to be of great advantage for us because of their ability to help in the clean up of
some contaminants. If phenol is present in low concentrations in water, Fontinalis antipyretica can
decompose this phenol up to 32–43%, and Platyhypnidium ripariodes can decompose them up to
20–27%. According to Mouvet et al. (1985), Cinclidotus danubicus, an aquatic bryophyte, is a good
292 � Bryophyta
accumulator of polychlorinated biphenyls. Mouvet et al. (1986) proved further that Fontinalis and
some other aquatic mosses “could be used to monitor both cadmium and polychlorinated biphenyls
because of their high accumulation ability.”
Takaki (1977) proved that “in some cases, pollution actually increases the cover of bryophytes.”
Bowden et al. (1994) studied that water bodies rich in phosphorus show an extensive growth of
Hygrohypnum alpestre and H. ochraceum.
BRYOPHYTES AS CLEANING AGENTS OF TOXIC WASTE 23.12

Some bryophytes, e.g. Sphagnum, show great promise to function as cleaning agents of toxic waste.
Rozmej and Kwiatkowski (1976) have opined that “even microorganisms have been cleaned up by
Sphagnum, perhaps due to the antibiotic properties of peat”. Calymperes delessertii, a moss, is also an
efficient adsorbent for dyes. According to Crum (1988), peat moss (Sphagnum) is “especially effective
at removing nitrogen (96%) and phosphorus (97%) applied from sewage or eutrophic river water”. Peat
has also been used to clean wastewater containing oil. In some countries, like Finland and Canada, peat
is also used “as a filter agent for oily waste in vegetable oil factories” (Ruel et al., 1977). According to
Viraraghavan and Tanjore (1994), the “highly toxic pentachlorophenol (PCP) is readily adsorbed by
Sphagnum peat.” Such an adsorption is irreversible, and this makes Sphagnum peat “an effective and
inexpensive means of removing such toxicants.” In some countries, including Poland, Sphagnum is
now being widely sold for reclaiming strip-mined land. Some scientists have also suggested that peat is
“a possible material for filtering water for reuse in space travel” (Crum, 1988).
1. Give a brief account of “bryophytes as indicators of environmental condi ons and pollu on”
in about 500 words.
2. Discuss “ecological uses of bryophytes” in about 500 words.
3. Do pollutants in bryophytes inhibit sexual reproduc on?
4. Giving suitable examples, give an account of indicator species of bryophytes in about 200
words.
5. “Bryophytes control erosion”. Comment in about 100 words.
6. Write a brief scien fic note on bryometer or moss bag.
7. An apparatus used to monitor changes in the atmosphere of bryophytes is named _____
8. Why do mosses and liverworts suit well as bioindicators? Write at least three such
characteris cs of bryophytes.
9. Explain in brief the role of “bryophytes as bioindicators of heavy metals in air pollu on.”
10. What are copper mosses?
11. Comment on “bryophytes as indicators of radioac vity”.
12. Explain the following in about 100 words each:
(a) Bryophytes as cleaning agents of toxic waste
(b) Bryophytes as indicators in aqua c habitats
24
Biologically Active
Compounds from
Bryophytes
WHAT ARE BIOLOGICALLY ACTIVE COMPOUNDS? 24.1
A great variety of terpenoids, aromatic compounds and acetogenins have been isolated from
bryophytes. Many of these have characteristic scents, pungency and bitterness, and show extraordinary
bioactivities as well as medicinal properties. These terpenoids, aromatic compounds and acetogenins
are included under biologically active compounds.1
SOME BIOLOGICALLY ACTIVE COMPOUNDS

EXTRACTED FROM BRYOPHYTES 24.2
Amongst bryophytes, members of Hepaticae contain cellular oil bodies which are extracted easily with
organic solvents, while members of Musci and Anthocerotae do not possess such cellular oil bodies.
A large number of bryophytes have been used as medicinal plants in China and many other countries,
specially to cure burns, bruises, external wounds, dermatitis, etc. Most of the Hepaticae (liverworts)
“contain mainly lipophilic mono-, sesqui-, and diterpenoids, aromatic compounds (bibenzyls, bis-
bibenzyls, benzoates, cinnamates, long-chain alkyl phenols, naphthalenes, phthalides, isocoumarins)
and acetogenins which constitute the oil bodies” (Asakawa, 2007). Y. Asakawa of Japan made elaborate
studies on this aspect and has isolated over 400 new compounds from bryophytes (Asakawa, 1990,
1993, 1995, 2007).
Some mosses and liverworts of medicinal importance and possessing biological activity are listed
in Table 24.1.
1.
For details, consult article of Yoshinori Asakawa (2007) of Japan, published in Pure Applied Chem. Vol. 79,
No. 4, pp. 557–580.
294 � Bryophyta
Table 24.1 Biological ac vity and effects of some liverworts and mosses of medicinal importance
NO. PLANTS BIOLOGICAL ACTIVITY AND EFFECTS

(A) Liverworts (Hepaticae)
1. Conocephalum conicum Antifungal, antimicrobial, antipyretic and antidotal activity; used
to cure burns, cuts, fractures, snakebites, and swollen tissues.
2. Frullania tamarisci Antiseptic activity.
3. Marchantia polymorpha Antihepatic, antidotal, antipyretic, diuretic activity; used to cure
burns, scalds, open wounds, poisonous snakebites, cuts, etc.
(B) Mosses (Musci)
1. Bryum argenteum Antidotal, antipyretic, antirhinitic activity.
2. Ditrichium pallidum For convulsions in infants.
3. Funaria hygrometrica For pulmonary tuberculosis, bruises, dermatomycosis.
4. Haplocladium catillatum Antidotal and antipyretic activities; pharyngitis, mastitis,
uropathy, pneumonia, tymphanitis and urocystitis.
5. Plagiopus oederi As sedative; used for epilepsy and cardiopathy.
6. Polytrichum commune Antidotal and antipyretic; cuts, hemostasis, bleeding from
gingivae, tuberculosis of lungs.
7. Rhodobryum roseum As sedative, used also for cardiopathy and neurasthenia.
CHARACTERISTICS OF BIOLOGICALLY ACTIVE

COMPOUNDS ISOLATED FROM BRYOPHYTES 24.3
Yoshinori Asakawa (2007) listed the following characteristics of the biologically active compounds
(terpenoids and aromatic compounds) isolated from liverworts:
1. Characteristic scents
2. Pungency and bitterness
3. Dermatitis
4. Cytotoxic, anti-HIV, and DNA polymerase b-inhibitory
5. Antifungal and antimicrobial activity
6. Insect antifeedant activity, mortality and nematocidal activity
7. Superoxide anion radical release inhibitory activity
8. Enzymes and NO production inhibitory activity
9. Piscidal and plant growth inhibitory activity
10. Neurotrophic activity
11. Muscle-relaxing activity
12. Inhibitory activity against osteoporosis and allergy
13. Cardiotonic and vasopressin antagonist activity
14. Anti-obesity activity
15. Synthesis of bioactive compounds from constituents of liverworts
Some brief aspects of all these characteristics of the biologically active compounds isolated from
bryophytes are discussed below.
Biologically Ac ve Compounds from Bryophytes � 295
CHARACTERISTIC SCENTS 24.4

Several simple aromatic compounds or volatile terpenoids are emitted by many members of Hepaticae.
These compounds are responsible for several scents, which are sweet-woody, sweet-mossy, carrot-
like, mushroomy or seaweed-like. Some such Hepaticae members, possessing characteristic odours or
scents (Asakawa, 2007), are listed below:
1. Asterella species produce indole- or skatole-like odour.
2. Conocephalum conicum produces strong mushroomy or camphor-like odour.
3. Frullania davulica produces mossy odour.
4. Jungermannia obovata produces carrot-like odour.
5. Lophozia bicrenata produces pleasant cedar-oil-like odour.
6. Pellia endiviifolia produces dried seaweed-like odour.
7. Porella gracillima produces woody-earthy odour.
8. Takakia lepidozioides produces mixed smell of cinnamon and burnt wheat powder.
9. Targionia hypophylla produces sweet terpentine-like odour.
Biocyclohumulenone
Tamariscol
O Grimaldone
H
O
HO
H
C
A
B
Fig. 24.1A-C Characteris c odorants from some liverworts. A, Biocyclohumulenone;

B, Tamariscol; C, Grimaldone (a er Asakawa, 2007)
Mushroom-like smell of almost all liverworts is due to the compound OCT-1-en-3-01 and its acetate.
Bicyclohumulenone (Fig. 24.1A) has been isolated from Plagiochila sciophila while tamariscol
(Fig. 24.1B) has been isolated from Frullania davulica. Both these compounds of commenre are used
as “perfumes as such or as perfume components of powdery floral-, oriental bouquet-, fantastic chypre-,
fancy violet-, and white rose-types in various cosmetics” (Asakawa, 2007). The “sweet terpentine-like
odour of Targionia hypophylla is due to a mixture of cis- and trans- pionocarveyl acetates” (Asakawa,
2007). The compound grimaldone (Fig. 24.1 C) is responsible for the strong sweet-mossy smell of
Mannia fragrans.
PUNGENCY AND BITTERNESS 24.5

Very intense pungent and bitter substances, which show interesting biological activities, are produced
by some liverworts. Porella vernicosa and some other species of this liverwort contain potent pungent
substances. Its strong, hot and pungent taste is due to the compound (–) -polygodial (Fig. 24.2A).
296 � Bryophyta
Several compounds possessing an intense pungent taste have been isolated by Asakawa (2007) and his
co-workers from Plagiochila, Pellia and Trichocoleopsis. Sacculatal and 1 b-hydroxysacculatal (Fig.
24.2B) have been reported from the ether extract of Pellia endiviifolia. When one chews a whole plant
of Plagiochila asplenioides and some of its other species, one feels a potent hot taste slowly. This is due
to the compound plagiochiline-A (Fig. 24.2C).
CHO R CHO
CHO CHO
(A)
R=H (B)
R=OH (C)
Fig. 24.2 Characteris c pungent and bi er substance e.g. polygodial (A), sacculatal and
1b- hydroxysacculatal (B) and plagiochiline (C) from some liverworts
DERMATITIS 24.6
Very intense allergenic contact dermatitis is caused by some liverworts including Frullania species.
The substances responsible for inducing allergy include sesquiterpene lactones, (+) – frullanolide (Fig.
24.3A) and (–) –frullanolide (Fig. 24.3B). Allergenic contact dermatitis is also caused by Marchantia
polymorpha and Metzgeria furcata.
Fig. 24.3 Some allergy-inducing compounds isolated from Frullania sp.; A, (+) - Frullanolide; B, (–) - Frullanolide
CYTOTOXIC, ANTI HIV 1, AND DNA

POLYMERASE b INHIBITORY ACTIVITY 24.7
Cytotoxic activity against lymphocytic leukemia is shown by some Hepaticae including Bazzania
pompeana, Porella japonica, and Radula perrottetii. Marsupellone and acetoxymarsupellone isolated
from Marsupella emarginata also showed cytotoxicity against certain types of lymphocytic leukemia.
Riccardin-A and Riccardin-B from Riccardia multifida inhibited KB cells. Plagiochila species also
show similar characteristics. Marchantia polymorpha, which can cause allergenic contact dermatitis,
shows inhibitory activity against Gram-positive bacteria. It also has diuretic activity. Marchantin-A
isolated from M. polymorpha, also shows cytotoxicity against lymphocytic leukemia.
ANTIFUNGAL AND ANTIMICROBIAL ACTIVITY 24.8

Antimicrobial activity is shown by several liverworts including the species of Bazzania, Dumortiera,
Marchantia, Plagiochila and Radula. Liverworts, which show antifungal activity include Lunularia
cruciata, Marchantia polymorpha and Plagiochila vernicosa, besides many others. Antibacterial
activity against Bacillus cereus, Cryptococcus neoformans, Escherichia coli, Pseudomonas aeruginosa
and Salmonella typhimurium is shown by the compound Marchantin-A obtained from several species of
Marchantia, including M. polymorpha, M. plicata and M. tosana (Asakawa, 2007). Marchantin-A also
has antifungal activities against fungi such as Alternaria kikuchiana, Aspergillus fumigatus, A. niger,
Candida albicans, Penicillium chrysogenus and Trichophyton rubrum. Strong antibacterial activity is
shown against Streptococcus mutans (dental caries) by the compound sacculatal (Fig. 24.2) isolated
from Pellia endeviifolia.
INSECT ANTIFEEDANT, MORTALITY AND

NEMATOCIDAL ACTIVITY 24.9
Species of Plagiochila contain plagiochiline-A (Fig. 24.2 C), which is a strong insect antifeedant,
specially against the worm Spodoptera exempta. Plagiochiline-A also shows nematocidal activity
against the nematode Caenorphabdiitis elegans. Succulatal (Fig. 24.2 B), a compound obtained from
several bryophytes (e.g. Trichocoleopsis), kills tick species (e.g. Panonychus citri). Insect antifeedant
activity is also shown by several species belonging to Chiloscyphus, Gymnocolea and Plagiochila.
SUPEROXIDE ANION RADICAL RELEASE

INHIBITORY ACTIVITY 24.10
Angiopathies are serious heart-related problems, such as cardiac infarction and arterial sclerosis. These
may be caused by excess superoxide anion radical (O –2) in organisms. Plagiochilal-B (Fig. 24.4A)
obtained from species of Plagiochila, norpinguisone methyl ether from Porella elegantula and radulali-
n–K (Fig. 24.4 B) from Radula javanica are examples of some of the compounds which inhibit the
release of superoxide anion radical from guinea pigs.
298 � Bryophyta
O
OH
H
OHC
HOOC
OHC
H O
A B
Plagiochilal-B Radulanin-K
Fig. 24.4 A, Plagiochilal-B; B, Radunalin-K
ENZYMES AND NITRIC OXIDE PRODUCTION

Production of several enzymes is inhibited by the activity of several compounds produced by many
liverworts. The enzymes include 5-lipoxygenase, hyaluronidase, cyclooxygenase, etc., and compounds
include marchantin-A, marchantin-B, marchantin-E from Marchantia, radulanin-H from Radula, and
riccardin-C from Riccardia. Lunularic acid, found in almost all liverworts, as a minor component, has
antihyaluronidase activity. In several liverworts “overproduction of NO is involved in inflammatory
response-induced tissue injury and the formation of carcinogenic N-nitrosamines. Large amounts of
NO were expressed and generated by iduced iNOS on stimulation of endotoxins or cytotoxins involved
in pathological response. Thus, inhibition of iNOS is very important to control inflammatory disease”
(Asakawa, 2007).
PESTICIDAL AND PLANT GROWTH

Porella vernicosa and several other species of this liverwort produce (–)-polygodial (Fig. 24.2), which is
a pungent and very strong pesticide. Sacculatal (Fig. 24.2), obtained from some other species of Porella
(e.g. P. endiviifolia) also possesses similar pesticidal properties. Both polygodial and sacculatal can
kill certain kinds of fishes within two hours. “Almost all crude extracts from liverworts which contain
bitter or pungent substances show phytotoxic activity”. Polygodial also promotes root elongation in
rice dramatically.
NEUROTROPHIC ACTIVITY 24.13

Compounds like mastigophorenes A, B and D (Fig. 24.5), obtained from Mastigophora diclados
show neurotrophic properties in fetal rat hemisphere according to Fukuyama and Asakawa (1991).
Acceleration of neurotic sprouting of neuronal cell culture of fetal rat cerebral hemisphere is also seen
in the compounds obtained from Plagiochila fruticosa.
Fig. 24.5 Mas gophorene A, B and D obtained from Mas gophora diclados
MUSCLE RELAXING ACTIVITY 24.14

Marchantin-A, obtained from Marchantia polymorpha and some other species (e.g. M. paleacea)
of this genus, is a pharmacologically important compound because of its muscle-relaxing activity.
Marchantin-A, along with its trimethyl ether, shows surprising muscle-relaxing activity.
INHIBITORY ACTIVITY AGAINST OSTEOPOROSIS

AND ALLERGY 24.15
Matsunaga et al. (1993) and Katsunuma (1997) have established the role of some liverworts (e.g.
Porella japonica in inhibitory activity against osteoporosis and allergy. Scientists in Japan are working
to develop chemopreventive drugs for these diseases from some liverworts.
300 � Bryophyta
CARDIOTONIC AND VASOPRESSIN

ANTAGONIST ACTIVITY 24.16
According to Asakawa (1990), marchantin-A, obtained from Marchantia polymorpha, shows cardiotonic
activity because it increases coronary blood flow. Prenyl bibenzyl, a compound obtained from Radula
perrottetii, shows vasopressin antagonist activity.
ANTIOBESITY ACTIVITY 24.17

Compounds, such as riccardin-C and ricccardin-F, have been isolated from some bryophytes (e.g.
Reboulia hemispherica and Blasia pusilla). They function as liver-X-receptors (LXR) μ- antagonist
and liver-X-receptor b-antagonist. Riccardin-C also increases cholesterol efflux from some cells, and
is used in the development of some drugs having antiobesity activity.
SYNTHESIS OF SOME BIOACTIVE COMPOUNDS FROM

CONSTITUENTS OF LIVERWORTS 24.18
Mastigophora diclados produces herbertane dimers, such as mastigophorene-A and mastigophorene-B
(Fig. 24.5) which possess neurotrophic activity. Ptychantin-A (Fig. 24.6), produced from the liverwort
Ptychantus striatus of family Lejeuneaceae has properties of treating several disorders such as blood
pressure, glaucoma, congestive heart failure and bronchial asthma, according to detailed studies of
Hagihara et al. (2003, 2006).
Fig. 24.6 Ptychan n-A produced from Ptychantus striatus

1. What are biologically ac ve compounds? Write a note on some biologically ac ve compounds

in about 200 words.
2. Write an essay on biologically ac ve compounds found in bryophytes.
3. Y Asakawa of Japan isolated approximately how many biologically ac ve compounds from
bryophytes?
4. Make a list of at least seven characteris cs of the biologically ac ve compounds isolated from
bryophytes. Elaborate at least one of them in about 100 words.
5. Give a brief account of biological ac vi es and effects of Marchan a polymorpha, Funaria
hygrometrica and Polytrichum commune.
6. Explain in brief the role of bryophytes in trea ng derma s, fungal and other microbial
infec ons.
7. Name a bryophyte from which a compound obtained shows cytotoxicity against lymphocy c
leukemia.
8. Compounds obtained from which bryophyte have an obesity ac vity?
25
Bryophytes
and Their Role
in Carbon and
Nitrogen Cycling
ROLE OF PLANTS IN REGULATING
BIOGEOCHEMICAL CYCLES 25.1
Plants play critical and definite roles in regulating biogeochemical cycles. Growth of plants controls
the exchange of gases that support life in our biosphere. Growth of plants also affects soil development.
Plants are our primary producers, and due to this, they also influence the distribution of energy for
higher trophic levels.
DO BRYOPHYTES HAVE THE SAME ROLE AS VASCULAR PLANTS IN

REGULATING BIOGEOCHEMICAL CYCLES? 25.2
Bryophytes do not have the same role as vascular plants in regulating biogeochemical cycles. It is
because of their unique physiology and ecology in which bryophytes differ from vascular plants in
influencing cycles of elements (e.g. C, N), energy and water. For example, an effective water-relation
system is present in bryophytes. Two phenomena, which allow bryophytes to tolerate longer periods
of water stress than vascular plants and also allow them to recover quickly with rehydration, are
poikilohydry and desiccation tolerance. (Poikilohydry is a phenomenon of lacking structures or
mechanisms to regulate water loss and, hence, having water content determined rapidly by the water
potential of the environment, as in algae, bryophytes).
Due to poorly-developed conduction systems, water and solutes are taken up over the entire plant
surface in bryophytes. Characters such as lack of (i) effective gametophyte stomata, and (ii) effective
cuticle in many species of bryophytes, allow free exchange of gases and solutions across cell surfaces.
Because of such characters, bryophytes often serve as effective traps for water and nutrients. In
Bryophytes and Their Role in Carbon and Nitrogen Cycling � 303
comparison to vascular plants, this quality of bryophytes also makes them more sensitive to atmospheric
chemical deposition.
HOW DO BRYOPHYTES INFLUENCE

ECOSYSTEM FUNCTIONS? 25.3
According to Turetsky (2003), bryophytes influence ecosystem functions due to the characters related
to their specific physiology and life history by
1. Producing organic matter,
2. Stabilising soils or debris,
3. Trapping sediment and water, and
4. Providing food and habitat for algae, fungi, invertebrates and amphibians.
There exist several mechanisms by which bryophytes influence carbon (C) and nitrogen (N) cycles,
and some such mechanisms are detailed in this chapter.
MAJOR ROLES OF BRYOPHYTES IN C AND N CYCLING 25.4

Mentioned below are some of the major roles of bryophytes in carbon and nitrogen cycling, as also
pointed out by M R Turetsky (2003) in an article published in the Bryologist:
Generally, the bryophytes
1. Fix C and N from atmospheric pools,
2. Reduce N availability for vascular plants and microbes,
3. Release dissolved compounds that are immobilised by soil microbes and lost via runoff, and
4. Transform C and N into recalcitrant organic matter.
ABILITY OF BRYOPHYTES TO INFLUENCE

LOCAL SOIL CLIMATES 25.5
Bryophytes have definite impact on local soil climates. They influence local soil climates by
1. Increasing soil moistures,
2. Decreasing soil temperature, and
3. Changing the density of soil organic matter.
The roles and influences mentioned above under Articles 25.4 and 25.5 confirm that bryophytes
influence the share of “C and N ecosystem inputs, and indirectly influence the rate at which these
elements are lost from ecosystems through litter decay, fire, and herbivory” (Fig. 25.1; Turetsky,
2003).
304 � Bryophyta
Fig. 25.1 Scheme summarizing the influences of bryophytes on carbon and nitrogen cycling in terrestrial
ecosystems, including posi ve (+) and nega ve (–) influences on the microbial ac vity, fire and
runoff (adapted from Turetsky, 2003)
ROLE OF BRYOPHYTES IN CARBON CYCLING 25.6

25.6.1 Carbon Fixa on by Bryophytes
Houghton et al. (2001) opined that one of the largest flow or continuous change in the global C cycle is
between atmospheric CO2 and land vegetation. The carbon derived from autotrophic fixation through
the process of photosynthesis makes a very large amount and comprises about half of all organic matter.
It has been established that bryophytes directly influence the flow of C into “ecosystems through their
metabolism and growth rates. While estimates of bryophyte biomass have been characterised in various
ecosystems, annual accumulations of C in plant material are more useful for studies of the C cycle.”
Carbon gains through net primary production (NPP) can be determined through net exchange of CO2
(Tuba et al., 1998) or annual production of biomass (Swanson and Flanagan, 2001; Ilyashuk, 2002).
(NPP represents the difference between gross primary production and primary respiratory losses. Gross
primary production represents total amount of organic matter produced per unit time).
In comparison to tropical bryophytes, the production rates of temperate and polar bryophytes have
been relatively well studied. From this point of view, worth-mentioning studies have been made on
Polytrichum alpestre and Chorisodontium aciphyllum by Fenton (1980), and on Sphagnum by Schofield
(2001) and Vitt et al. (2003).
Clarke et al. (1998) reported growth of tropical epiphytic bryophytes ranging from 122–203 g
biomass m–2 yr–1. The growth of the canopy and gap species may be higher than that of understorey
bryophytes, as estimated by workers like Losch et al. (1994) end Zotz et al. (1997).
Sand-Jenson et al. (1999) opined that aquatic bryophytes often dominate vegetation in lakes, and
bryophyte NPP can exceed that of algae. Bryophytes exploited from this point of view by various
bryologists include Sphagnum (Lannergren and Ovstedal, 1983), Jungermannia vulcanicola (Miyazaki
and Satake, 1985) and Fontinalis hypoides (Ilyashuck, 2002).
25.6.2 Control on Growth of Bryophytes

Mentioned below are some of the factors which internally control the photosynthetic efficiency in
bryophytes:
1. Rate of photosystem electron transport.
2. Morphology of plant, in general, and ‘leaf’, in particular.
3. Environmental factors such as light and water. In species of Sphagnum, water limitation
“decreases resistance to C uptake” according to Rice (2000).
4. Availability of nutrients also strongly influence C fixation by many bryophytes, especially
terrestrial species.
5. Growth of temperate species of Sphagnum is promoted by increasing photoperiod length.
6. Increase in the radiation flow also helps in promoting the growth of temperate species of
Sphagnum.
7. Factors, such as declining photoperiod length and declining night temperature, induce
dormancy in mosses.
8. Both high and low water contents limit C uptake in bryophytes. Photosynthetic enzyme activity
is inhibited in conditions of low water availability.
9. Effect of water stresses varies among different species of bryophytes. Higher water content
was most stressful for Leucobryum antillarum, while growth of many tropical bryophytes
(e.g. Frullania mirabilis, Phyllogonium fulgens) was more limited by low water availability
according to Zotz et al. (1997).
10. Growth of bryophytes is also influenced substantially by “altered moisture availability”
(Turetsky, 2003).
11. According to Vitt et al. (2003) and many other workers, “as N is thought to limit plant growth
in many terrestrial systems, bryophyte productivity commonly increases following N or
combined N and phosphorus (P) additions”.
12. Arscott et al. (1998) have correlated high phosphorus inputs to high bryophyte productivity.
306 � Bryophyta
25.6.3 Impact of Dissolved Organic Carbon

According to Moore and Dalva (2001), “carbon can be leached from plant biomass as dissolved organic
carbon”. A major part of this dissolved organic carbon can be readily utilised by microbes, and its
remaining part “can be comprised of highly refractory compounds with complex structures” (Turetsky,
2003). In this way, the dissolved organic carbon can either be “utilised by microbes or can be lost from
terrestrial ecosystems via runoff”. It is this terrestrial dissolved organic carbon which is transported to
aquatic bodies like lakes, estuaries or oceans, where it represents an important transfer of energy and
carbon from terrestrial to aquatic systems. This dissolved organic carbon is very important and has the
ability to change the production in greenhouse gases, trace metal speciation, acid-base chemistry and
N and P availability according to workers like Gergel et al. (1999).
In general, it has been proved that plants influence the leaching of dissolved organic carbon through
production of soluble organic compounds. As far as bryophytes are concerned, the “species common to
boreal peatlands differ in water-soluble carbohydrates, phenolics, and soluble non-polar compounds”,
according to Turetsky (2003). In Frullania atrata, the concentrations of soluble sugars and polyols
comprised up to 17% of dry weight in the upper canopy of forests, while in Phyllogonium fulgens, it is
equivalent to only 6% of dry weight in lower canopy, according to Coxson et al. (1992). In mosses, the
“greater amounts of soluble proteins and carbohydrates are associated with higher metabolic activity”
(Pakarinen and Vitt, 1974).
According to Coxson et al. (1992), the release of soluble sugars from epiphytic bryophytes in tropical
forests is equivalent to 122 kg ha–1yr–1. Carleton and Read (1991) estimated leakage of carbohydrates
from Pleurozium schreberi after the extended summers and concluded that “these carbohydrate-rich
leachates are capable of supporting mycorrhizal fungal growth.”
According to Charman et al. (1999) and Chasar et al. (2000), some amount of dissolved organic
carbon in peatlands reaches downwards into the peat and is used by microbes. Fraser et al. (2001)
opined that export of dissolved oxygen-carbon can be a significant loss of carbon from the terrestrial
ecosystem.
In spite of all these above-mentioned studies, Turetsky (2003) mentioned that “more research is
needed to understand the long-term fate of bryophyte-derived dissolved oxygen-carbon in terrestrial
and aquatic ecosystems”.
25.6.4 Decomposi on and Implica ons for Soil Carbon

After the death of plants, carbon in their organic matter is degraded through the action of bacteria and
fungi. The rate of decomposition is affected also by various factors such as temperature, moisture,
etc. According to Merrifield and Ingham (1998), bryophytes “may influence microbial activity by
providing microhabitat for invertebrates”. Tsuneda et al. (2001) opined that “bryophytes can also
harbour microfungi that decompose organic carbon”. Eckstein (2000) suggested that “bryophytes
also influence decay by reducing soil temperature and/or increasing soil moisture. Since bryophytes
have low thermal conductance, they can increase water availability through external capillary action”
(Turetsky, 2003).
Litter with poor organic-matter quality is produced by bryophytes. At the same time, bryophytes
are also not able to synthesize lignin. Due to the absence of lignin, bryophyte litter would decay more
rapidly than vascular material of higher plants. Organic matter produced by bryophytes generally
decomposes very slowly. In Sphagnum, the slow rate of decomposition may be because of low N
concentrations. Decay is also inhibited in bryophytes due to the presence of large concentrations of
phenolics and nonpolar compounds. Verhoeven and Liefveld (1997) have identified polyphenolic
networks in mosses resembling vascular lignins and tannins. These compounds can mask cellulose and
also inhibit microbial breakdown, and can also make cell walls impenetrable to hyphae of fungi.
Verhoeven and Toth (1995) have worded on antimicrobial properties of bryophytes, such as
Sphagnum. Basile et al. (1999) have shown that flavonoids isolated from mosses have antibacterial
properties. Banerjee and Sen (1979) have focused in detail on the antibiotic activity of 52 species of
bryophytes.
Rate of decomposition, however, differs widely among different species of bryophytes, e.g. decay
process is very slow in species of Sphagnum than other mosses (Belyea, 1996).
ROLE OF BRYOPHYTES IN NITROGEN CYCLING 25.7

25.7.1 Biological Fixa on of Nitrogen
The largest global pool of nitrogen is the atmosphere. Plants need nitrogen for the production
of chlorophyll and RUBISCO (Ribulose 1, 5- biphosphate carboxylase oxygense) and also for the
construction of proteins and nucleic acids. This nitrogen comes from biological fixation, atmospheric
deposition and weathering. In many ecosystems, the largest source of nitrogen is by biological fixation.
Many prokaryotic microorganisms (e.g. Cyanobacteria) use the nitrogenase enzyme to break the triple
bonds of atmospheric nitrogen and fix it into more soluble forms. Large amount of energy is spent in
this nitrogen-fixation process. Symbiosis occurs between nitrogen-fixing Cyanobacteria and hosts (e.g.
algae, fungi, bryophytes and many vascular plants). In this way, bryophytes also influence biological
nitrogen fixation. These Cyanobacteria may be epiphytic or endophytic, e.g. Nostoc in the thalli of
Anthoceros. Meeks et al. (1983) proved that “ammonium (NH +4) is the initial product of N fixation by
symbiotic Nostoc associated with the hornwort Anthoceros punctatus. A small portion of N2-derived
ammonium (~10%) is assimilated by Nostoc and the remaining product is transferred efficiently to host
tissue and utilised as amino acids.”
Vitousek (1994) concluded that “liverworts are important to N fixation in Hawaiian forests.” Sheridan
(1991) estimated that “Hapalosiphon flexosus–Sphagnum erythrocolyx associations contribute about
400 mg N m–2 yr–1 on a tropical volcanic dome.” Basilier (1980) reported that “N fixation in coniferous
forests occurred only with Sphagnum plants.”
Devey and Marchant (1983) and some other workers investigated that availability of moisture
“appears to be important to N fixation”, specially in plants like Andreaea, Ditrichum strictum and
Jamesoniella colorata. Temperature is also a strong control on N fixation in bryophytes of polar regions
(Davey and Marchant, 1983). Sheridan (1991) observed that “wind or volcanic disturbance increases
fixation by tropical Hapalosiphon flexosus–Sphagnum erythrocolyx.
25.7.2 Assimila on of Nitrogen by Bryophytes

Bryophytes have access to inorganic N in the form of ammonia and nitrates and/or organic N, according
to Lipson and Nasholm (2001). Nitrates (NO3–) and nitrites (NO2–) are reduced after assimilation.
According to Brown (1992), “bryophytes generally assimilate NH4+ more readily than NO3–.” Miyazaki
and Satake (1985) concluded that “liverworts such as Jungermannia vulcanicola and Scapania undulata
used NH4+ as their major N source”. According to Rudolph et al. (1993), low pH usually inhibits NO3–
308 � Bryophyta
assimilation. Glime (1992), however, opined that “acid-tolerant aquatic species such as Sphagnum
and Drepanocladus rely heavily on NH4+ for N requirements.” Brown (1992) mentioned further that
“bryophytes are able to take up organic N such as amino acids or dipeptides.” Studies of Kielland
(1997) illustrate that organic N is an important source of nitrogen for bryophytes such as Cetraria
richardsonii and Sphagnum rubellum. Turetsky (2003) opined that “bryophytes generally are very
efficient in assimilating N, and appear to rely mainly on atmospheric deposition.” Tracer studies of
Li and Vitt (1997) and Lamontagne et al. (2000) using 15N highlight the mechanisms of N uptake and
retention by plants. Svensson (1995) concluded that “bryophytes are competitive scavengers of N and
reduce N availability for higher plants.” Species of Sphagnum are extremely efficient in capturing N
because the entire plant is able to absorb nutrients according to Rudolph et al. (1993). Turetsky (2003)
stated that “epiphytic bryophytes play an important role in N cycling within canopies, though less is
known about their N-use efficiency.”
25.7.3 Loading of Nitrogen and Bryophyte Assimila on

Loading of nitrogen can generally change plant productivity and patterns of N retention. It “can also
influence community composition by favouring nitrophilous species” (Turetsky, 2003). Nordin and
Gunnarsson (2000) reported that ammonium nitrate (NH4+NO3–) applications increased amino acid
concentrations in Sphagnum but NH4+ alone decreased organic acid concentrations in other bryophytes
according to Soares and Pearson (1997).
Pitcairn et al. (2002) opined that bryophytes are often used in determining programmes as indicators
of N pollution in ecosystems. Several studies (Penuelas and Filella, 2001) are available which prove
that mosses have been used for temporal assessment of N deposition.
25.7.4 Transforma ons and Losses of Nitrogen

Uptake of nitrogen by plants and microorganisms is very rapid, and due to this, the majority of soil
N occurs in organic forms. Nitrogen from organic matter is released by activity of microbes. This
loss of organic N from ecosystems is reduced by processes such as (i) decreasing decomposition, (ii)
herbivory, and (iii) combustion. Studies suggest that bryophytes may enhance N availability for higher
plants if N slowly released from the breakdown of bryophyte litter can be taken up more efficiently.
It has also been proved that dissolved organic nitrogen is released from organic matter of the plants
through processes like (i) microbial breakdown, and (ii) leaching. Willimas et al. (1999) studied this
aspect of dissolved organic nitrogen in Sphagnum recurvum and S. capillifolium. In case the dissolved
organic nitrogen is not immediately immobilised, it can be exported from terrestrial ecosystems via
runoff.
WHAT MORE CAN BE DONE IN UNDERSTANDING ROLE OF

BRYOPHYTES IN C AND N CYCLING? 25.8
Some of the knowledge gaps still inhibiting our understanding of the role of bryophytes in C and N
cycling are listed below:
1. Major researches in this aspect have only been done on mosses. Much more is still to be done
on liverworts and hornworts.
2. A majority of the tropical bryophytes have not been investigated so far from the point of view
of role of bryophytes in C and N cycling.
3. The role of bryophytes in biogeochemical cycling should be investigated not as a single
functional group but it should also be investigated along with plants of other groups.
4. Due emphasis should be given to understand the mechanisms controlling bryophyte chemistry
while studying the role of bryophytes in C and N cycling.
5. Usually, bryophyte material is of poor litter quality. It should also, therefore, be investigated
in detail that how this poor quality litter influences C and N cycling.
6. Bryophyte productivity should also be measured properly prior to estimating their role of C
and N cycling.
7. It should also be estimated thoroughly that “how will changes in species diversity influence
ecosystem processes or controls on ecosystem processes” (Turetsky, 2003).
8. Whether genetic diversity of bryophytes is also an important factor in C and N cycling should
also be studied in detail.
9. Tolerance levels of various species of bryophytes to N deposition should also be investigated
while studying their role in C and N cycling.
10. It should also be observed whether species disappear in response to N pollution.
11. Whether deposition of N changes bryophyte biochemistry should also be worked out while
studying the role of bryophytes in C and N cycling.
12. Will global warming influence bryophyte chemistry should also be studied properly?
1. Write an essay on bryophytes and their role in carbon and nitrogen cycling.
2. Do bryophytes have the same role as vascular plants in regula ng biogeochemical cycles?
Comment only in about 100 words.
3. What is poikilohydry?
4. How do bryophytes influence func ons of the ecosystem?
5. Describe the role of bryophytes in carbon cycling.
6. Write at least five factors which internally control the photosynthe c efficiency in
bryophytes.
7. Discuss in brief the role of bryophytes in the nitrogen cycle.
8. What do you mean by biological fixa on of nitrogen, with par cular reference to
bryophytes?
9. What is the full form of RUBISCO?
26
Evolution of the
Gametophyte in
Bryophytes
VIEWS ON THE EVOLUTION OF THE
GAMETOPHYTE IN BRYOPHYTES 26.1
Two broad types of views have been put forward by bryologists to explain the process of the evolution of
gametophyte in bryophytes. Some bryologists are of the opinion that the simple thallus of Marchantiales is
the result of retrogressive evolution while the other group of bryologists believe that the Marchantiaceous
thallus is simple because of progressive evolution.
Some aspects of the mechanisms of both retrogressive as well as progressive types of evolution are
briefly discussed below:
MECHANISM OF RETROGRESSIVE EVOLUTION 26.2

This theory has been supported by Wettstein (1903–1908), Goebel (1906), Kashyap (1919), Evans (1939),
Mehra (1953, 1957), Zimmermann (1966) and Udar (1970). These workers believe that the primitive
gametophyte was an erect leafy shoot of Bryopsida showing radial symmetry. A reduction in the leaf
development in such a primitive bryophytic member of Bryopsida resulted in the evolution of the
dorsiventral thalli in acrogynous Jungermanniales, Marchantiales and Anthocerotales.
According to Wettstein (1903–1908), a hypothetical primitive ancestor may be traced in the leafy
gametophytes of Calobryales and several true mosses, such as Fontinalis, Polytrichum and Bryum. The
possible approach to such a hypothetical primitive ancestor may also be traced in the sexual branches
of some prostrate forms of thalloid and leafy Hepaticopsida. Ultimately, the dorsiventral thalli of several
Hepaticopsida and Anthoceropsida developed from such ancestors by the mechanism of retrogressive
evolution, i.e. progressive reduction.
Evans (1939) was an active supporter of the mechanism of retrogressive evolution and had provided
several examples to explain the evolution of the thalloid bryophytes from the erect, radial leafy shoots of
foliose members. As the dorsiventral bilaterality of the leafy shoot progressed, a gradual reduction in the
size of the ventral leaves took place, as seen in genera such as Lepidozia and Lejeunea. The process of
Evolu on of the Gametophyte in Bryophytes � 311
reduction in the size of the leaves ultimately reached such an extent that they appeared in the form of
mucilaginous papillae, as in Plagiochila, and finally they disappeared in Radula.
Along with this character of gradual reduction and ultimate disappearance of ventral leaves, there was
also a flattening of the axis as well as a gradual decrease in the size of lateral leaves either in the form of
a few cells or slime papillae. Sometimes even their complete disappearance was noticed as in Pellia,
Marchantiales and Anthocerotales. Some intermediate stages of the members, such as Pellia, may be seen
in genera, such as Zoopsis and Schiffneria—in which the leaves are more or less reduced and the axis is flat.
However, in several such examples (e.g. Marchantia), the reproductive shoots (i.e. antheridiophores and
archegoniophores) remain radial, erect and somewhat leafy in nature, and this indicates that the thalloid
state is one of secondary origin. On the basis of his studies of genus Monoselenium of Marchantiales, Goebel
(1906) had also favoured the mechanism of retrogressive evolution of gametophytes in bryophytes.
Shiv Ram Kashyap (1919), the noted Indian bryologist, made extensive studies on Indian liverworts,
noted several examples of reduction series in genera belonging to Marchantiales, and became a strong
supporter of the mechanism of retrogressive evolution. Three main points of this reduction series suggested
by Kashyap (1919) are given below:
1. The Loss of Assimilatory Filaments in Air Chambers The stages of the gradual reduction
in the assimilatory filaments can be observed in four different genera of bryophytes as under:
1. In Preissia quadrata (Fig. 26.1A), a species occurring on moist soil, all the structures of the
higher forms (i.e. air chambers, air pores and assimilatory filaments) are present.
2. In Conocephalum conicum and other genera growing in more moist places, the assimilatory
filaments are short (Fig. 26.1B) and pointed.
3. In Weisnerella denudata (Fig. 26.1C), growing near or under water, the air chambers contain
only papillate cells.
4. In Dumortiera (Fig. 26.1D), growing under water, the air chambers are either absent completely or
present only at the growing points. The air pores are rudimentary, and the assimilatory filaments
are absent.
Fig. 26.1A-D Gradual reduc on of the assimilatory filaments in Marchan ales (A, Preissia quadrata;
B, Conocephalum conicum; C, Wiesnerella denudata; D, Dumor era hirsuta)
312 � Bryophyta
2. Simplifica on of Air Pores The air pores are barrel-shaped on both thallus as well as on discs of
antheridiophores and archegoniophores in Marchantia; they are barrel-shaped on the discs but simple
on the thallus in Conocephalum; they are simple on both, thallus as well as discs in Exormotheca;
and the air pores are not well-developed at all in Riccia.
3. Gradual Shi ing of the Erect Branches bearing Sex Organs to the Dorsal Position due
to the Continued Growth of the Thallus and Gradual Elimina on of the Stalk In
Marchantia, the two sex organs, i.e. antheridia and archegonia, develop on special erect branches called
‘antheridiophores’ and ‘archegoniophores’, respectively, and both these are usually terminal in position.
In Plagiochasma articulatum and some other genera, the stalk is initially terminal in position but soon
becomes dorsal because of the continued growth of the thallus. In Corsinia, the reduction trend reaches a step
further as the female receptacle becomes sessile due to the gradual elimination of the stalk.
On the basis of his studies on a fossil Hepaticopsida—Naiadita, Harris (1939) also opined that “at a
certain stage, the primitive Hepaticae had an erect leafy shoot with spirally arranged leaves as in moss.”
Genetical studies of Burgeff (1943) also supported the theory of retrogressive evolution. Burgeff evolved
numerous mutations of Marchantia and correlated several of them with the phyletic reduction series.
Mehra (1957) proposed the condensation theory, according to which the Marchantiaceous thallus has
been derived from foliose Jungermanniales through the processes of compaction and condensation. Mehra
suggested that the type of the thallus represented by Stephensoniella brevipedunculata and Asterella
reticulata is the most primitive in Marchantiales and is very near to the ancestral type. “This, in turn, seems
to be developed from foliose forms by the processes of compaction, condensation and fusion of leaves”
(Mehra, 1957).
THEORY OF PROGRESSIVE EVOLUTION 26.3

This theory has been supported by bryologists such as Cavers (1910), Campbell (1918, 1936, 1940),
Smith (1955), etc. According to this theory, the primitive gametophyte was a dorsiventral and prostrate
thallus which was simple in its morphological as well as anatomical characters.
Fig. 26.2A-C Thalli of Sphaerocarpos s pitatus (A), Riccardia indica (B), and Metzgeria pubescens (C)
Cavers (1910), the main supporter of this theory, opined that the thallus of the present-day genus
Sphaerocarpos (Fig. 26.2A) resembles the simplest primitive gametophyte. However, Campbell (1918,
1936, 1940) believed that the nearest approach to the simplest primitive gametophyte is to be seen in the
thalli of the living genera Riccardia and Metzgeria (Fig. 26.2B,C). Therefore, the simple and ‘’primitive
type of thalli according to this theory are of Sphaerocarpos, Riccardia and Metzgeria. The evolutionary
advance in such simple thalli progressed in two different lines:
Evolu on of the Gametophyte in Bryophytes � 313
1. One line of the evolutionary advance resulted in the formation of the thalli of Marchantiales
(e.g. Marchantia) by the gradual formation of structures, such as epidermis with air pores, air
chambers, assimilatory filaments, and separate reproductive branches, i.e. antheridiophores and
archegoniophores.
2. The second line of evolutionary advance resulted in the formation of the gametophytes of
Jungermanniales and Calobryales, in which the internal structures remained simple, due to the
absence of air pores and air chambers, but the external structures of the gametophyte elaborated
finally into a foliose or leafy shoot.
1. Give an account of the evolu on of the gametophyte in bryophytes in about 500 words.
2. Descrive the mechanism of retrogressive evolu on of gametophyte in bryophytes.
3. Who have been the two main Indian advocates of mechanism of retrogressive evolu on of
gametophytes in bryophytes?
4. Enumerate three main points of reduc on series in evolu on of gametophyte suggested by
Shiv Ram Kashyap.
5. Describe the theory of progressive evolu on of gametophyte in bryophytes in about 200
words.
27
Origin and
Evolution of
Sporophyte in
Bryophytes
WHAT IS EVOLUTION OF THE SPOROPHYTE
IN BRYOPHYTES? 27.1
The zygote is the first cell of the sporophytic generation. It divides and redivides to form the sporophyte,
which is never an independent body in bryophytes. It is always dependent on the gametophyte, either
completely or partially. The function of the sporophyte is the production of spores.
Two different theories have been put forward by the bryologists to explain the evolution of the
sporophyte in bryophytes. These are (i) theory of the progressive sterilization of the potentially
sporogenous tissue, and (ii) theory of progressive simplification or reduction theory.
THEORY OF PROGRESSIVE STERILIZATION

OF POTENTIALLY SPOROGENOUS TISSUE 27.2
This theory, also called the theory of sterilization, has been proposed by Bower (1908), and supported
by Cavers (1910) and Campbell (1918, 1940). According to this theory, Riccia possesses the simplest
and the most primitive sporophyte, and from such a sporophyte of Riccia have evolved more advanced
sporophytes, through the process of progressive sterilization of potentially sporogenous tissue. A
definite series of such an evolution of the sporophyte from Riccia may be traced in the sporophytes of
Sphaerocarpos, Targionia, Marchantia, Pellia, Anthoceros and Funaria. A brief discussion of all these
stages from Riccia to Funaria follows:
1. First Stage The first stage is seen in Riccia (Figs. 27.1, 7.10) in which the zygote divides several
times to form a multicellular diploid structure, of which the outermost layer develops into a sterile
jacket while all the central mass remains sporogenous in nature. Each cell of this sporogenous tissue
Origin and Evolu on of Sporophyte in Bryophytes � 315
divides meiotically to form four haploid spores. Thus, the only sterile part of this simple sporophyte is
the sterile jacket, while the entire remaining tissue is fertile. Also, there is no differentiation of organs,
such as foot, seta and capsule, and, therefore, the sporophyte of Riccia is the simplest and the most
primitive (Bower, 1908).
Fig. 27.1 Sporophyte of Riccia
2. Second Stage In Riccia crystallina and Oxymitra, the sporophyte, of course, consists of a
single-layered jacket enclosing the central mass of the sporogenous tissue like that of other species of
Riccia discussed above. But some potential spore mother cells remain unable to form the spores and
form the sterile or abortive nutritive cells. Thus, the sterile tissue in the sporophyte is a little more in
amount than that of the first stage, i.e. other species of Riccia.
3. Third Stage In both Corsinia and Sphaerocarpos (Figs. 27.2, 5.2), more amount of potentially
sporogenous tissue becomes sterilised than that of Riccia crystallina and Oxymitra. In Corsinia, the
whole of the basal part of the sporophyte gets sterilised to form the foot, made up of a few cells. And in
Sphaerocarpos (Fig. 27.2), the basal part of the sporophyte is sterilised into a bulbous foot and a narrow
two-cells broad seta. In both Corsinia and Sphaerocarpos, a single-layered sterile jacket surrounds the
sporogenous tissue in the capsule. A few sterile nurse cells are also present in both along with the
fertile spores. The nurse cells, however, lack the characteristic spiral thickenings of the elaters.
Capsule
Spore
tetrad
Nurse cell
Seta
Foot
Fig. 27.2 Sporophyte of Sphaerocarpos s pitatus

316 � Bryophyta
4. Fourth Stage In Targionia (Fig. 27.3), the sporophyte contains still larger amount of the sterile
region in the form of a bulbous foot, narrow seta, single-layered jacket of the capsule, and several
elaters, containing spiral thickenings. Sterile elaters present along with the fertile spores, constitute
about half of the number of the sporogenous cells of the capsule.
Jacket
Capsule
Spore tetrad
Elaters
Seta
Foot
Fig. 27.3 Sporophyte of Targionia hypophylla
5. Fi h Stage In Marchantia (Figs. 27.4, 7.32), the sterile tissue in the sporophyte is slightly
more in amount than that of Targionia, and consists of a broad and bulbous foot, elongated and more
developed seta, single-layered sterile jacket of the capsule, a few sterile cells at the apex of the capsule
in the form of a small apical cap, and a large number of long elaters containing the spiral thickenings,
along with the fertile spores in the capsule.
Fig. 27.4 Sporophyte of Marchan a polymorpha
6. Sixth Stage In Pellia (Figs. 27.5, 4.16F), Riccardia and other Jungermanniales, still larger
percentage of the part of the total sporophyte is sterilised. The sterilised tissue consists of an elaborate
foot, well-developed seta, two to many-layered sterile jacket of the capsule, several elaters, and a mass
of sterile cells in the form of elaterophore. The elaterophore is either basal (Pellia) or at the apex
(Riccardia) of the capsule. Only a small percentage of the sporogenous tissue actually remains fertile
and forms spores.
Fig. 27.5 Sporophyte of Pellia epiphylla
7. Seventh Stage Highly reduced sporogenous tissue is present in Anthoceros (Figs. 27.6, 8.7
I-M) on account of further sterilisation. The sterile tissue of the sporophyte consists of massive foot,
small meristematic zone, 4- to 6-layered wall of the capsule, 4- to 16-celled thick columella and a large
number of pseudoelaters. Further, the sporophyte of Anthoceros shows a greater degree of independence
since its capsule wall possesses a well-defined epidermis, several stomata and chlorophyll-containing
cells.
Fig. 27.6 Sporophyte of Anthoceros
8. Eighth Stage The process of the progressive sterilisation of potentially sporogenous tissue
reaches at its peak in some higher members of Bryopsida, e.g. Funaria (Figs. 27.7, 10.22 B) and
Polytrichum. In Funaria, the sterile tissue of the sporophyte consists of foot, seta, entire region of the
318 � Bryophyta
apophysis of capsule, many-layered capsule wall, spore sac wall, columella, peristome and operculum.
The only fertile region is in the form of two spore sacs in the theca region of the capsule. Due to the
presence of stomata, very long seta, well-developed capsule wall, and large number of chlorophyll-
containing cells in the capsule, the sporophyte in these bryophytes shows still greater degree of
independence. According to the theory of progressive sterilisation of potentially sporogenous tissue
of Bower (1908), the sporophyte of Funaria, therefore, is the most advanced and highly evolved.
Fig. 27.7 Sporophyte of Funaria hygrometrica
THEORY OF PROGRESSIVE SIMPLIFICATION 27.3

This theory, also called the reduction theory, has been proposed by Church (1919) and supported
by Kashyap (1919), Goebel (1930) and Evans (1939). According to these bryologists, the simplest
sporophyte of Riccia is not a primitive type of the sporophyte. On the other hand, they believe that
the Riccia sporophyte is a reduced type evolved by a process of descending or regressive evolution or
“progressive simplification”.
According to the reduction theory of Church (1919), the ancestral sporophyte of Bryophyta was
an erect, foliose (leafy), and independent shoot like that of mosses. During the descending course of
evolution, such an erect leafy sporophyte passed through the following changes:
(i) It became attached to the gametophyte on the permanent basis, (ii) its leaves were lost due to the
prolonged isolation and desiccation, (iii) because of its greater dependence on the gametophyte, the
intercellular spaces of its photosynthetic system disappeared, (iv) in the later stages its assimilatory
tissue also became reduced, (v) the stomata of its epidermis first reduced into simple pores and became
functionless, as in Sphagnum, and then they completely disappeared, as in Marchantia and Riccia.
Church (1919), followed by Goebel (1930) and Evans (1939), hence, believed that the complex and
well-developed sporophytes of Funaria and Anthoceros, having intercellular spaces in their assimilatory
tissue and stomata in their epidermis, are primitive and nearer to the ancestral type. On the other hand,
the sporophytes of Pellia, Marchantia, Targionia and Sphaerocarpos are reduced and simplified, and
this series of reduction reached at its peak in the sporophyte of Riccia.
1. Give an illustrated account of evolu on of sporophyte in bryophytes.

2. What is the first cell of a sporophy c genera on?
3. In bryophytes, the sporophyte is never an ……………. body.
4. In bryophytes, the sporophyte is always dependent on ………….., completely or par ally.
5. The main func on of sporophyte in bryophytes is the produc on of ………………. .
6. What are the two major theories which have been put forward to explain evolu on of
sporophyte in bryophytes?
7. Explain in detail the “theory of sterilisa on” given to explain origin and evolu on of sporophyte
in bryophytes.
8. According to the theory of sterilisa on, which of the bryophy c genus possesses simplest
and most primi ve sporophyte?
9. In the undermen oned series of evolu on of sporophyte, what is missing?
Riccia Æ Sphaerocarpos Æ Targionia Æ Marchan a Æ Pellia Æ Anthoceros Æ ---------.
10. Draw labelled diagrams of the sporophytes of
(a) Anthoceros
(b) Riccia
(c) Pellia
(d) Funaria
11. With reference to the evolu on of sporophyte in bryophytes, what is the theory of progressive
simplifica on?
12. Theory of progressive simplifica on is also called ------------.
28
Conduction in
Bryophytes
HOW DO BRYOPHYTES MAINTAIN POOR INTERNAL
CONDUCTING STRANDS? 28.1
Being terrestrial, most bryophytes lack well-organised internal conducting strands. This deficiency of
bryophytes is, however, overcome by some characteristics, of which some are mentioned below:
1. Usually, bryophytes grow in moist shady places, with abundant humidity.
2. The gametophytic plant body of bryophytes absorbs water from all over the surface.
3. Most of the thalloid bryophytes lie prostrate, i.e. parallel to the ground. Due to this, the area of
contact, with the substrate or ground, increases.
4. The gametophytic plant body of such bryophytes, which are not prostrate but erect, is generally
small. This reduces the distance through which the water has to actually move inside the plant
body.
5. Plants show many adaptations which enhance or encourage external conduction.
EXTERNAL CONDUCTION AND ITS SIGNIFICANCE 28.2
28.2.1 External Conduc on in Gametophytes

Several structures on the gametophytic plant body of bryophytes are responsible for external conduction
of water, e.g. closely-placed branches on the main axis or leaves on the stem (e.g. Polytrichum) help in
external conduction. Leaves possessing sheathing leaf bases (e.g. Polytrichum) also facilitate external
conduction. Some mosses (e.g. members of Dawsoniaceae and Polytrichaceae) have lamellae on the
leaves, and these lamellae help in external conduction. Extensive growth of paraphyllia on the stem of
some mosses help in conduction of water.
Conduc on in Bryophytes � 321
Fig. 28.1 Ver cal sec on of thallus and basal part of archegoniophore of Fimbriaria bleumeana showing
rhizoidal groove (a er Bowen, 1935)
Some mosses show growth of their gametophytes in very close tufts, forming compact cushion-like
structures with effective capillaries between the gametophytes, as in pin-cushion moss (Leucobryum
glaucum). External conduction takes place through these capillaries. Several appendages develop on
the stem of some mosses. These appendages produce capillary channels. Water moves through these
channels in such gametophytes.
Pegged rhizoids and overlapping scales of many members of Marchantiales (e.g. Marchantia) also
help in external conduction. Such structures make channels for effective water transport.
Rhizoidal grooves in the archegoniophore of Marchantia and Fimbriaria (Fig. 28.1) also serve the
purpose of external conduction. Efficient capillary channels are also formed by twining together of
rhizoids in some bryophytes, e.g. Pogonatum.
28.2.2 External Conduc on in Sporophytes

External conduction in sporophytes in bryophytes is quite different from their gametophytes because
of some definite reasons, as under:
1. Appendages are absent in sporophytes, and therefore, there are no capillaries for external
conduction.
2. Mature sporophytes of mosses have seta and capsule bearing a thick cutinised epidermis
followed by a thick-walled hypodermis. Both of them prevent direct absorption of water
from their surrounding atmosphere. Some external absorption, however, occurs when the
sporophyte is very young.
3. In liverworts, the seta of the sporophytes elongates only at maturity followed by quick dispersal
of spores. So, the conduction is not needed at all.
28.2.3 Significance of External Conduc on

Because of the absence of a well-developed conducting tissue in the gametophytes of most of the
bryophytes, the external conduction of water by capillary action is of definite significance in these
plants. The role played by external conduction of water mainly depends on the morphology and anatomy
of the particular species of bryophyte. Environmental conditions (e.g. relative humidity) also plays a
definite role in external conduction. Bopp and Stehl (1957) have experimentally proved that external
conduction is more efficient than internal conduction in Funaria hygrometrica. Deloire et al. (1979)
determined experimentally the rate of external conduction in some more mosses and also in liverworts
322 � Bryophyta
by employing fluorescent dye. Eosin due was used by Clee (1937) to study external conduction in
Plagiochila and Pellia. In Conocephalum, McConaha (1939) proved experimentally that “conduction
of water along the entire length of the thallus takes about 20 to 30 seconds”. In the later years, the same
author studied the rate of external conduction in some more bryophytes including Reboulia (0.5 mm/s),
Lunularia (0.5 mm/s) and Preissia (1 mm/s). In Polytrichum, it has been proved by Magdefrau (1936)
that “external conduction is sufficient to maintain turgidity at 90% relative humidity, but at 70%, both
internal and external conduction are necessary”.
INTERNAL CONDUCTION AND ITS SIGNIFICANCE 28.3

Cells Involved in Internal Conduc on
In the gametophytic plant body, all the cells, to some extent, are capable of some degree of conduction.
But the function of conduction in the cells is related to several characteristics including (i) increase in
the length and breadth of cells, (ii) development of structures like plasmodesmata, small pores, etc.,
and (iii) thickening of the side walls of the cells, which provide support to the plant body. Wide variety
in the structure of cells, with reference to the conduction of water, is shown by bryophytes. It has,
however, been observed that poorly-developed conducting strands are seen when plants are very young,
and also when they grow under high humidity in their surroundings.
(a) Cells Involved in Conduc on in Liverworts and Hornworts Ordinary parenchyma cells in the
ventral region of the thallus are mainly responsible for conduction in a majority of the liverworts and
hornworts. According to Proskauer (1961), prominent primary pit fields are present in the walls of
these cells, as in Dendroceros crispus (Fig. 28.2). In the centre of the stem of many leafy liverworts and
also in the midrib region of thalloid bryophytes, some cells are elongated and possess plasmodesmata
in their walls. Such cells are called conducting parenchyma, as in Conocephalum. Empty and dead
cells form clear conducting strands in several liverworts including Hymenophyton, Haplomitrium and
Takakia (Fig. 28.3). According to Hebant (1977), the end walls of these mature dead and empty cells in
Takakia have small plasmodesmata-derived pores (Fig. 28.4 A, B), More advanced type of conducting
cells are present in the midrib region in Pallavicinia lyellii (Fig. 28.5).
Fig. 28.2 Cells from the thallus of Dendroceros crispus showing well-developed primary pit fields
Fig. 28.3 Transverse sec on of a part of the stem of Takakia showing true conduc ng strand
(a er Hebant, 1977)
Fig. 28.4 A, Diagramma c representa on of the conduc ng cells from the stem of Takakia
showing small plasmodesmata-derived pores in their end walls; B, Some part of
‘A’ (highly enlarged) showing details of end wall (a er Hebant, 1977)
324 � Bryophyta
Fig. 28.5 Cross sec on (a part) of the midrib por on of the thallus of Pallavicinia lyellii
showing conduc ng strand (a er Smith, 1966)
(b) Cells Involved in Conduc on in Mosses Conducting parenchyma, hydroids, leptoids and stereids
are some major types of cells involved in conduction in mosses.
(i) Conducting parenchyma occurs commonly in mosses in their stem, leaves and sporophyte.
Plasmodesmata and pits derived from them are present in these cells.
(ii) Hydroids are the water-conducting cells of mosses, e.g. Polytrichum (Fig. 28.6). They are
found in stem and leaf of gametophyte and seta of sporophyte. The hydroids collectively
constitute the hydrom.
(iii) Leptoids are the food-conducting cells found in some mosses e.g. Polytrichum (Fig. 28.6).
Along with the accompanying parenchyma, leptoids are collectively called leptom.
(iv) The midrib region of the leaves and axis of the stem in some mosses contain some thick-
walled elongated cells along with hydroids and leptoids. These are known as stereids, as in
Polytrichum (Fig. 28.6). The stereids are collectively called stereome.
Fig. 28.6 TS Rhizome of Polytrichum commune showing hydroids, leptoids and stereids
According to Hebant (1977), hydroids and leptoids are present in both gametophytes as well as
sporophytes (seta) in Polytrichum. In Funaria, hydroids are present in gametophyte (stem) as well as
sporophyte (seta) whereas leptoids are present in seta and absent in the stem. In Buxbaumia, hydroids
are present in the seta only, while they are absent in gametophytes (stem). Leptoids are absent in both
stem as well as seta of Buxbaumia. Both hydroids and leptoids are absent in both gametophytes (stem)
as well as sporophyte (seta) in Orthotrichum (Hebant, 1977). Members of Polytrichales are, therefore,
the only bryophytes, the gametophytic generation of which possess leptoids (i.e. food-conducting
cells).
ANATOMY OF MIDRIB AND LEAF TRACES 28.4

28.4.1 Midrib
The following three types of cells (Fig. 28.7) are found in the most complex midribs of leaves of
mosses:
Fig. 28.7 Ver cal sec on of the leaf of Mnium undulatum showing three types of cells
1. Deuters These are the large-sized, living cells which conduct food. They have plasmodesmata
in their end walls.
326 � Bryophyta
2. Hydroids These are the cells responsible for conduction of water.
3. Stereids These are thick-walled cells which serve as supporting cells in the midrib.
28.4.2 Leaf Traces

Leaf traces, if present, are not as complex as the midribs. The midrib may or may not be connected to
the central strands of the axis. The leaf traces may be true or false. If a leaf trace joins with the central
strand, it is called true. But if it does not come in contact with the central strand, it is called a false leaf
trace.
SIMILARITIES AND DIFFERENCES BETWEEN HYDROIDS OF

BRYOPHYTES AND TRACHEARY ELEMENTS OF PRIMITIVE
VASCULAR PLANTS 28.5
28.5.1 Similari es
Hydroids of bryophytes resemble tracheary elements of primitive vascular plants in the main events of
their development. Some such events are:
1. Increase in length and other parameters of dimensions
2. Obliqueness of end walls
3. Thick lateral walls
4. Degeneration of their protoplasts, making the cells empty and finally dead
5. End walls getting partially hydrolysed,
All these characteristics make hydroids the preferential pathways for water conduction, as is also
seen in primitive vascular plants.
28.5.2 Differences
1. Secondary thickenings in the form of spirals, rings or reticulum are absent in hydroids while
present in tracheary elements of primitive vascular plants.
2. Nature of perforations in the end walls of hydroids is also different from that of tracheary
elements of primitive vascular plants.
SIMILARITIES AND DIFFERENCES BETWEEN

LEPTOIDS OF POLYTRICHALES AND SIEVE ELEMENTS
OF VASCULAR PLANTS 28.6
28.6.1 Similari es
Typical leptoids are sieve-element-like cells. Some features, in which leptoids of Polytrichaceae
resemble closely with the sieve elements of vascular plants, are listed below:
1. Lengthening and broadening of their extremities

2. Oblique placement of their end walls
3. Presence of well-developed plasmodesmata in their end walls
4. Thickened lateral walls
5. Presence of at least some amount of callose in their end walls
6. Presence of refractive spherules in the leptoids similar to that of phloem of pteridophytes
7. Similar to that of phloem, leptoids start translocation of food even when they are still not fully
mature.
28.6.2 Differences
Leptoids differ from the phloem of vascular plants in some of the following characteristics:
1. Organisation of pores in their end walls
2. Persistence of degenerated nucleus
3. Absence of p-protein.
INTERNAL CONDUCTION OF WATER BY GAMETOPHYTE 28.7

Internal conduction of water in mosses (e.g. Mnium and Polytrichum) takes place by hydroids in the
stem, and the same was demonstrated by Haberlandt (1886) using eosin and lithium sulphate solutions.
Hebant (1974) also confirmed the same. The rate of conduction is 120 cm/h in Mnium and 200 cm/h
in Polytrichum.
True conducting strands are rarely present in liverworts. Smith (1966) demonstrated that “eosin
solution travelled up to 1.5 cm in 4 to 5 minutes in the conducting strand of Symphyogyna circinata”.
The K-fluorescein solution, however, travelled slightly rapidly in Takakia lepidozioides according to
Hebant (1972). Some studies suggest that water travels along the cell walls of all tissues of gametophytes
of mosses.
INTERNAL CONDUCTION OF WATER BY SPOROPHYTE 28.8

Sporophytes of bryophytes lack capillaries on their surface, and they also do not show appreciable direct
absorption of water. Therefore, the entire water required by the capsule of the sporophyte is supplied
internally. Seta of the sporophyte has well-developed conducting tissues, whether the conducting strands
are present or absent in the gametophyte. Bryologists have demonstrated the internal conduction in the
seta of Polytrichum, Funaria and some more mosses using eosin solution and fluorescent dyes.
CONDUCTION OF ORGANIC COMPOUNDS

IN BRYOPHYTES 28.9
With the help of 14C-bicarbonate (a labelled compound), it has been shown by Eschrich and Steiner
(1967) that in the stem of Polytrichum, the organic compounds are conducted at the rate of 32 cm/h.
Conduction of organic compounds (assimilates, exogenously applied sucrose and ionic solutes such as
328 � Bryophyta
lead and sulphate) in the stem of Polytrichum takes place through leptom, according to Trachtenberg and
Zamski (1978). In the seta of the sporophyte of Polytrichum, translocation of organic compounds takes
place at the rate of 50 cm/h, as demonstrated by Eschrich (1975) using 14C-sucrose. It has now been
finally proved that in Polytrichum, the organic compounds travel through leptoids in both gametophytic
and sporophytic generations.
Bopp and Knoop (1974) demonstrated the internal conduction of organic substances in the protonema
of mosses while Rota and Maravolo (1975) studied conduction of labelled sucrose in the thalli
of Marchantia. Rose and Bopp (1983) demonstrated internal transport of indole-acetic-acid in
the rhizoids of Funaria hygrometrica.
1. Explain conduc on in bryophytes with the help of suitable diagrams.

2. Bryophytes have poorly developed conduc ng strands. How do they maintain their this
characteris c? Explain in about 100 words.
3. Describe external conduc on and its significance in bryophytes.
4. “Pin-cushion moss” is _____ _____.
5. In Marchan a, structures like overlapping scales and pegged rhizoids help in _____ _____.
6. Write a note on significance of external conduc on in bryophytes.
7. Give a brief descrip on of internal conduc on in bryophytes.
8. Which of the following are the cells involved in conduc on in mosses?
(a) Hydroids (b) Leptoids (c) Stereids (d) All of these.
9. With the help of only one sentence about each of them, differen ate between the following
with reference to their involvement in conduc on in mosses:
(a) Conduc ng parenchyma, (b) Hydroids, (c) Leptoids, (d) Stereides.
10. In Polytrichum, which of the following means of conduc on is/are found?
(a) Stereides, (b) Leptoids, (c) Hydroids, (d) All of these.
11. Name the only single order of bryophytes, the gametophy c genera on of which possess
leptoids.
12. What are deuters?
13. Leptoids are the _____ conduc ng cells of bryophytes.
14. Make a list of some similari es and differences between hydroids of bryophytes and tracheary
elements of primi ve vascular plants.
15. What are the similari es between leptoids of Polytrichales and sieve elements of vascular
plants?
16. Write a note on conduc on of organic compounds in bryophytes in about 100 words.
29
Elementary
Cytogenetics and
Cytotaxonomy
of Bryophytes
Cytogenetics has been such a field of research during the last few decades that a very vast literature is
available in the form of voluminous books and research publications. To give a glimpse of all aspects
of cytogenetics of bryophytes within few pages of a chapter is neither possible nor within the scope
of this book. Hence, only some selected preliminary aspects of this modern branch of bryophytes have
been dealt with here.
In the cytology, the aspects briefly discussed include (i) basic techniques, (ii) mitosis, (iii) meiosis,
(iv) heterochromatin, (v) chromosome number, and (v) extra chromosomes, including sex-associated
chromosomes.
In the genetical studies, some aspects discussed are (i) hybridization, and (ii) polyploidy.
Some details of cytotaxonomy have also been discussed at the end of the chapter.
Important initial contributions on the cytology of bryophytes have been made by Mahabale (1942),
Lowry (1954), Steere (1954), Bryan (1956), Vaarama (1956), Khanna (1960), Gangulee and Chatterjee
(1960), and Mehra and Khanna (1961).
SOME BASIC TECHNIQUES OF CYTOGENETICS 29.1

Rapid acetocarmine squash method is the first and most suitable karyological technique. It enabled
us to determine the chromosome number of several mosses. This technique was modified by Heitz
(1926). In place of orcein dyes (1.5 g of orcein mixed in 100 cc of 45% acetic acid in a flask), carmine
was largely replaced as a staining agent for studying bryological materials by Lowry (1948) and
Vaarama (1949). Cytology of many bryophytes has also been studied by Feulgen staining techniques
by workers, namely Darlington and La Cour (1950, Lewis (1957) and Vaarama (1964).
For studying fresh material of bryophytes, aceto-orcein stain gives good results. For squash
preparations of the archesporial tissue, the tissue from the capsule is squeezed out on a slide and
immersed in a drop of aceto-orcein. No prior fixation of the fresh material is required. The slide is
330 � Bryophyta
slightly heated and the pressure is applied with a needle. The squash preparation of the archesporial
tissue is now ready to study various stages of meiosis. The haploid chromosome number of the plant
can be determined by counting the meiotic bivalents by this technique. Lewis (1957) studied somatic
mitosis in liverworts and mosses by pretreatment of the material “with 8-hydroxyquinoline as a 0.002
molar aqueous solution”. Vaarama (1953) used the same technique for meiotic studies of several
mosses.
Temporary preparations are sealed by ringing the cover glass with rubber solution. For making
permanent preparations, the squash slides are passed through a slowly ascending alcohol series and
mounted in Euparol or other suitable mounting medium.
SOME STUDIES ON MITOSIS 29.2

C E Allen (1912) studied cell structures, growth and division in the antheridia of Polytrichum juniperum,
and later on worked on a chromosome difference correlated with sex differences in Sphaerocarpos (Allen,
1917). It was E Heitz (1928) who first reported heterochromosomes in Pellia epiphylla, a liverwort.
Heitz later on studied mitosis in liverworts and mosses. K Yan (1957), a Japanese bryologist, worked
on cytology of several Japanese mosses and equated sex chromosomes with heterochromosomes, but
the term “heterochromosome” is not a synonym of sex chromosome. A Vaarama made significant
contribution in cytology of bryophytes. He observed many heavily stained heteropycnotic bodies of
variable sizes within an interkinesis nucleus of Pleurozium schreberi (Vaarama, 1954). In the mitotic
chromosome of this bryophyte, the chromatids get separated from each other prior to anaphase even at
the locus of centromers. The chromatids remain attached together due to sticky matrix.
In some bryophytes, Heitz (1928) reported the presence of chromosomes and their segments “which
remained dense and deeply stained at stages (e.g. prophases) in contrast to the other chromosomes
which were in large part diffuse and lightly stained”. He described such chromosomes and their
parts as heteropycnotic. These chromosomes are made up of different material, which is known as
heterochromatin.
SOME STUDIES ON MEIOSIS 29.3

Since it is difficult to get the adequate preparations of the stages of meiotic prophase, the details of
the process of meiosis in most of the bryophytes is, therefore, difficult to study. In Grimmitaceae, a
family of mosses, meiosis has been studied by Vaarama (1949). The bivalent contains two chiasmata,
one in each chromosome arm. The chiasma is situated either in the proximal or in the distal part of the
chromosome arm. The frequency of the chiasma and also the number of interstitial chiasmata varies in
different species of mosses belonging to Grimmiaceae.
In almost all bryophytes studied so far, “the chromosome compliment contains a heterobivalent which
deviates from the autosome bivalent in its (i) larger size, (ii) asymmetry, and (iii) heterochromacity at
interkinesis, mitotic and meiotic prophase stages.” In several pleurocarpous mosses, the heteropycnotic
chromosome pair develops in a very specialised way during meiotic prophase. During pachytene stage,
the heteropycnotic chromosome, which lies on the surface of a vesicle-like body, disappears. In this
manner, a ring-shaped bivalent develops. This bivalent differs in appearance from the other bivalents.
Vaarama (1953) named this bivalent M-bivalent or special bivalent.”
Elementary Cytogene cs and Cytotaxonomy of Bryophytes � 331
Vaarama (1954) studied the structure and behaviour of meiotic bivalents in Hedwigia ciliata, a
moss. In diplotene-diakinesis stages of this moss, the chromatids of the special bivalent “are either
quite separate from each other or united pairwise or in chains or rings by terminal contacts which are
not chiasmata”. The chromatids are negatively heteropycnotic at diakinessis and metaphase-I stages.
The chromatid pairs are situated opposite each other at both sides of the metaphase plate. It appears that
the chromatids of special bivalent are quite independent in their function.
In Pleurozium schreberi, meiotic prophase starts with a stage exhibiting optically single strands,
according to Vaarama (1954). The chromatids of bivalent halves as well as chromosomes of anaphase-
II stage are “clearly double and the half-chromatids have matrices of their own” (Vaarama, 1954).
Frequent exchanges in the location of centromeric activity takes place in meiosis.
Meiosis in Sphagnum has been studied in detail by Sorsa (1955). He reported an unusual centromere
in Sphagnum, and also tripolar spindles in mosses. His researches have been published in Vol. I of
Hereditas. Sorsa (1955) described “the diploid chromosome number in Sphagnum sporophytes has 9
bivalents and 1 univalent while the species with tetraploid sporophytes (e.g. Sphagnum robustum and
S. squarrosum) were found to have 19 bivalents.” However, according to Bryan (1955) and Holmen
(1955), the “haploid or gametophytic number in Sphagnum is 19 plus the m-chromosomes and that the
diploid or sporophytic number is 38 (19 homologous pairs) plus the m-chromosomes”. Bryan (1955)
clearly observed 19 bivalents in addition to the m-chromosomes in the spore mother cell of Sphagnum
erythrocalyx.
Study of chromosomes with particular emphasis on meiosis in about 40 species of 16 genera of
American mosses was done by Steere et al. (1954). This has been published in Mem. Torrey Botanical
Club (Vol. 20). Several of these species show polyploidy. More than ten of these species possess the
so-called accessory chromosomes. Large-sized heteromorphic bivalent chromosomes were observed
in several species, and these were assumed by these workers as sex chromosomes. From these studies,
they concluded that “the cytological behaviour of bryophytes parallels that of phanerogams very
closely”.
Postmeiotic changes in Physcomitrella patens, Desmatodon randii and Funaria hygrometrica have
been studied by Ripetsky and Matasov (1975).
Six sub-stages of meiotic prophase in Ditrichum pallidum have been described by Brown and
Lemmon (1980), and their research has been published in Vol. 83 of Bryologist. Some aspects of these
stages are described below:
Stage 1 Just before meiosis, each sporocyte produces a thick layer of sporocyte wall. It consists
of fibrillar polysaccharides. This wall, present between archesporial cell wall and protoplast, survives
throughout the formation of spore. The archesporial cell wall dissolves and/or disintegrates, and due to
this, the sporocytes are released into the spore chamber. A large number of ribosomes are also present
in the cytoplasm.
Stage 2 Synaptinemal complexes start developing in this stage, which also show almost all features
of Stage 1.
Stage 3 Nuclei containing well-developed synaptinemal complexes start shifting towards sporo-
cyte periphery in this stage. They also “assume the acentric bouquet appearance”. Fine, tightly packed
unpaired chromosomes are present in the nucleus. Soon they show the typical pachytene structure.
332 � Bryophyta
Stage 4 Centrally-located pachytene nuclei are seen.

Stage 5 Diffused stage of nuclei with relaxed chromatin are seen. Cytoplasmic depressions pro-
duce lobes in tetrahedral arrangement. Several microtubules also now develop and also proliferate
around the plastid in each of the four lobes and finally ensheath the nucleus.
Stage 6 The nuclear membrane dissociates and the chromatin again condenses into chromosomes.
Soon, the appearance of kinetochore-microtubule attachment is seen.
Development of diploteine starts with the formation of chiasmata in the bivalents.
In yet another study, Brown and Lemon (1980) studied stages of meiosis in Ditrichum pallidum after
prophase. According to them, during metaphase-I in this species, “the bivalents are distributed along
the equator of an open spindle consisting of continuous microtubules. Microtubules are attached at
kinetochores”. The distribution of both plastids and mitochondria is associated with the lobing pattern
in the cytoplasm of metaphase-I sporocyte. Most notable is the positioning of one of the four plastids
into each of the cytoplasmic lobes. Meiosis- II occurs after a short intrameiotic interphase. Young
spores are separated within the tetrad. The spores are invariably arranged tetrahedrally.
HETEROCHROMATIN 29.4
Chromatin is the complex of proteins, DNA and small amounts of RNA of which chromosomes are
composed.
Heterochromatin is a condensed region of chromatin in the interphase nucleus that stains heavily
with basic dyes. Inactive nuclei contain large amounts of heterochromatin.
Euchromatin is an expanded region of chromatin in the interphase nucleus, which stains lightly
with basic dyes. Metabolically active nuclei show large amounts of euchromatin.
Chromatin of chromosomes is of definite help in their selective staining. It is specifically possible
during cell division when chromosomes are thick and condensed. Heitz (1928) was the first to see
and study heterochromatin in a bryophyte, while studying gametophytic cells of Pellia endivaefolia.
Presence of heterochromatin is of specific interest in sex-associated chromosomes. It distinguished sex-
associated chromosomes from otherwise similar chromosomes. According to the studies of Segawa
(1965) and Ono (1970), Y-chromosomes contain more heterochromatin than the X-chromosomes. Some
other details of heterochromatin are discussed under Article 29.6 (Sex-Associated Chromosomes).
CHROMOSOME NUMBER 29.5

Counting of chromosomes is of primary concern in the cytology of bryophytes. Several workers have
worked on this aspect of this fascinating group of plants. To name a few are Wilie (1957), Proskauer
(1958), Anderson (1962), Fulford (1965), Khanna (1965), Smith and Newton (1968), Moore (1970,
1977), Fritsch (1972, 1982), Smith (1978) and Newton (1977, 1984). Different groups of bryophytes
(Hepaticopsida, Anthoceropsida and Bryopsida) are quite different from each other in their chromosome
numbers.
29.5.1 Chromosome Number of Hepa copsida

Hepaticopsida generally show a uniformity in their chromosome numbers. Smith and Newton (1968)
and Smith (1978) reported n = 8 or 9 in liverworts. Calobryales and Sphaerocarpales have n = 9 or n
= 8 and n = 9, respectively. Marchantiales generally have the chromosome number as n = 9 or in its
multiple. In Riccia, it is n = 8, 16, 24 and 48. Jungermanniales have n = 9, while in Metzgeriales the
predominant number is n = 9 with a few deviations (e.g. n = 10 in Aneura and n = 20 in Riccardia).
29.5.2 Chromosome Number in Anthoceropsida

According to Proskauer (1958), the chromosome number in members of Anthocerotopsida is n = 5
while Berrie (1960) reported that it is n = 6 in species found in Japan. As mentioned earlier also,
members of Anthoceropsida, Hepaticopsida and Bryopsida have very different chromosome numbers
as reported by Newton (1984) while studying the frequency of chromosome numbers in this group
(Fig. 29.1).
Fig. 29.1 Frequency of chromosome number in all the three groups of bryophyta.
A, Anthoceropsida and Hepa copsida; B, Bryopsida (a er Newton, 1984)
29.5.3 Chromosome Number in Bryopsida

Frequency of chromosome number in Bryopsida, shown in Fig. 29.1B, shows that mosses are quite
variable in their chromosome number. However, many genera and families are considered haploid with
n = 6 and n = 7. Many of the moss genera with 10–14 chromosomes are considered basically diploid.
In Andreaeales, the chromosome number of n = 10 and n = 11 have been reported in three species
of Andreaea. The recent trend is to consider Takakia as a moss under the order Andreaeales, and Inoue
(1973) reported n = 4 and n = 5 for two species of Takakia, the lowest chromosome number in mosses.
334 � Bryophyta
Ulothrix, a green alga, also possesses n = 4, the lowest basic chromosome number in green algae, which
are thought to be the ancestors of bryophytes by scientists working on the phylogeny of this group.
A polyploid series of n = 9, 18, 27 and 54 has been reported in Physcomitrium pyriforme, a common
green-house moss of temperate regions. In Sphagnum, the most worked-out moss, chromosome
complement of n = 19 + 2 microchromosomes or 38 + 4 microchromosomes have been reported.
EXTRA CHROMOSOMES AND SEX ASSOCIATED

CHROMOSOMES 29.6
Chromosomes complements of bryophytes differ from that of other groups in several characteristics
including (i) size, (ii) degree of hetero-chromaticity, (iii) mode of pairing, and (iv) segregation in the
process of meiosis. Some specialised chromosomes are present in liverworts, e.g. sex chromosomes,
micro-chromosomes, and other heteropycnotic chromosomes.
29.6.1 Microchromosomes and Heteropycno c Chromosomes

Microchromosomes are dot-like or short rod-like structures, as in most Hepaticopsida. Extremely
small microchromosome is a small accessory chromosome found in members of Anthoceropsida and
Bryopsida.
Heteropycnotic chromosomes are very small and also called h-chromosomes, as in Takakia
lepidozioides, Calobryum rotundifolium and many mosses. A heteropycnotic chromosome is about
1/3 to ½ of the longest autosomes of the genus. According to Berrie (1968), an h-chromosome is of
“partly heteropycnotic nature combined with the fact that it is no longer than any other member of the
chromosome complement”.
29.6.2 Sex-associated Chromosomes

Allen (1919) was the first to discover sex chromosomes in Sphaerocarpos donnellii, a liverwort. Heitz
(1932) discovered them in Ceratodon purpureus, a moss. Since then, sex chromosomes have been
discovered and discussed in several bryophytes by many workers including Lorbeer (1934), Allen
(1945), Tatuno (1941), Lewis (1961), Vitt (1968), Khanna (1971), Berrie (1974), Newton (1979),
Fritsch (1982) and Kumar (1983).
The female specific X-chromosome in Sphaerocarpos donnellii is the largest member of the
complement (Fig. 29.2A), while the male specific Y-chromosome is the smallest (Fig. 29.2B). Lorbeer
(1934) reported in Lunularia cruciata and Oxymitra paleacea that Y-chromosome is larger in these
genera than X-chromosome, a rarely seen condition. In Pellia endivaefolia, Tatuno (1941) reported yet
another exception, where sex chromosomes are smallest members of the complement. Berrie (1974)
also reported the similar condition in Plagiochila praemorsa, a liverwort.
According to Lorbeer (1938, 1941), Sphaerocarpos shows strong evidence for X-and Y- as sex-
associated chromosomes. In this liverwort, the plants remain female even after the distal portion
of X-chromosome is deleted. Opposite to this, plants of Sphaerocarpos with “radiation-damaged
X-chromosome may be male in absence of Y-chromosome” according to Knapp (1936).
Fig. 29.2 A-B, Sex chromosomes in a complement of Sphaerocarpos donnellii;

largest one in A is X-chromosome from female plant, while the small one in B is
Y-chromosome from a male plant (a er Allen, 1919)
Most of the dioecious species of mosses possess sex-associated chromosomes. According to

Ramsay (1974), dioecious species of Macromitrium show some evidence for dimorphic X- and Y-
chromosomes. This characteristic is not, however, seen in monoecious species of Macromitrium. It
is also seen that monoecious plants are homosporous whereas dioecious plants possess two types
of spores, i.e. large and small. Larger spores develop into female plants while smaller spores give
rise to male plants. Heterochromatin of interphase nuclei also differs in female and male plants. Sex-
related nature of largest heterochromatic chromosome has also been confirmed by Newton (1971) in
Plagiomnium undulatum.
Mentioned below is a summary of sex-chromosome mechanism in liverworts, as proposed by Berrie
(1963):
(i) X/Y consisting of two large chromosomes X and Y;
(ii) x/y consisting of two small chromosomes x and y;
(iii) X/y consisting of a large X chromosome
and small y chromosome, as in Facultative heterochromatin
Sphaerocarpos; and Euchromatin Constitutive heterochromatin
(iv) X1, X2, Y mechanism in which the X
element is represented by X1 and X2
chromosomes, and a Y chromosome, as
in some species of Frullania.
Clear difference are observed in sex-
associated chromosomes in their structure and
morphological details in most of the investigated
species of bryophytes. According to Segawa
(1965) and Ono (1970), Y-chromosomes contain
more heterochromatin than X-chromosomes. Structural Morphological
Newton (1979) has presented a schematic Sex chromosomes
representation of the evolution of sex Fig. 29.3 Evolu on of sex chromosomes in Atrium
chromosomes in Atrium crispum on the basis of crispum (a er Newton, 1979)
336 � Bryophyta
their structure and morphological details. It possesses euchromatin, facultative heterochromatin and
constitutive heterochromatin (Fig. 29.3).
29.6.3 Nucleolar Chromosomes

Large heteropycnotic chromosomes, reported along with sex chromosomes or microchromosomes in
about half a dozen species of Marchantiales and Acrogynae, have been termed nucleolar chromosomes
by Berrie (1958, 1968). Nonheteropycnotic nuclear chromosome, attached terminally to the nucleolus,
have also been reported by several workers in Frullania lyellii, Pallavicinia lyellii and Riccardia pinguis.
According to Berrie (1963), “in the nucleolar chromosomes, the nucleoli are formed at telophase of cell
division on specific chromosomes”.
29.6.4 Accessory Chromosomes

Vaarama (1949) observed very small chromosomes in Grimmia muehlenbeckii, a moss. Since
they resembled the chromosomes of some flowering plants (e.g. Poa, Zea, etc.), they were named
as accessory chromosomes by Vaarama. Later on, accessory chromosomes were reported in many
other bryophytes, such as Orthotrichum tenellum, Dicranum majus, etc. Accessory chromosomes in
bryophytes are smaller than the normal chromosomes, and are also heterochromatic. Pairing takes
place only between two homologous accessory chromosomes. In all the studied species, the accessory
chromosomes are not situated in the normal plate either in the first or second metaphase. The accessory
chromosomes in most of the species are generally four in number. Rarely, their number may be only
two. Anderson and Bryan (1954) reported accessory chromosomes in 16 American species of mosses
and interestingly all these species were polyploids.
HYBRIDIZATION 29.7
A natural or artificial process that leads to the formation of a hybrid is known as hybridization.
A hybrid is a plant which results from the cross-fertilization of two different species, subspecies,
varieties, strains, etc.
Differing from higher plants, there are only few reports of natural hybridization in bryophytes. It was
Wettstein (1923) who made the first artificial cross between two mosses, viz. Funaria hygrometrica
and Physcomitrium pyriforme. In some later studies, Wettstein (1924, 1928) was successful in making
several interspecific crosses in species of mosses of Funariaceae and Bryaceae. Although hybrid
sporophytes are mostly sterile, in some cases viable spores are also produced by them. Based on
his studies, Wettstein (1928, 1932) concluded that the “cytoplasm as well as the chromosomes play
an important genetic role. Some characters of the gametophytic progeny are determined by genes,
the distribution being Mendelian; some are determined by the cytoplasm, the results being maternal
inheritance; some by the combined influence of genes and cytoplasm”.
Natural hybrids described in some bryophytic genera include Bryum, Dicranella, Ditrichum,
Funaria, Grimmia, Orthotrichum, Physcomitrella, Physcomitrium, Rhacomitrium, Tetraplodon and
Weissia.
Pettet (1964) studied some intergeneric hybrids of Funariaceae.
POLYPLOIDY 29.8
Cells, which have three or more sets of homologous chromosomes in their nuclei, are known as
polyploids, and the phenomenon of their formation is called polyploidy.
Polyploidy is not very common in bryophytes. However, it is seen in most of the members of
Marchantiales. According to Schuster (1966), about 15% of the liverworts show polyploidy. Hexaploid
forms of Dumortiera hirsuta and large-sized polyploid Targionia lorbeeriana with n = 27 have been
reported from Japan. In Riccia, polyploidy is seen only in those species which are capable of growing
in xeric conditions. Polyploidy is more common in mosses than liverworts. High degree of polyploidy
is observed in mosses belonging to Pottiaceae, Funariaceae and Bryaceae.
29.8.1 Autopolyploidy or Intraspecific Polyploidy

A polyploid species with all sets of chromosomes coming from the same species is called autopolyploid,
and this phenomenon is known as autopolyploidy or intraspecific polyploidy. Autopolyploidy has
been reported in several liverworts, including Asterella, Calypogea, Cephalozia, Dumortiera, Nardia,
Pellia, Riccardia, Riccia and Targionia. Amongst mosses, autopolyploidy has been reported in many
genera including Atrichum, Bryum, Distichium, Funaria, Mnium, Pohlia, Sphagnum and Weissia.
Several cytologists recognise several polyploid, taxa as independent species (Newton, 1990).
In some mosses, the autopolyploids are morphologically different but in others they are
morphologically same. To mention a few examples, “Sphagnum subsecundum (n = 19 + 2m) and S.
auriculatum (n =38 + 4m) differ only in the size of their organs” according to Hill (1975). However,
according to Smith and Hill (1968), in Funaria hygrometrica (n = 28, 56), Atrium undulatum (n = 7,
14, 21) and Physcomitrium pyriforme (n = 26, 52), “there are no differences between autopolyploids
or cytotypes”.
29.8.2 Allopolyploidy or Interspecific Polyploidy

A polyploid species with sets of chromosomes from two or more different species is called
allopolyploid, and this phenomenon is called allopolyploidy. This can be the result of hybridization
between species.
Allopolyploidy is not well recorded in liverworts. Most of the mosses also do not show allopolyploidy.
Wettstein (1932), however, recorded 28 interspecific hybrids in mosses. Out of these 28 hybrids, “24
are between species which are monoecious”. In comparison to monoecious species, a low frequency of
hybrids is seen between species which are dioecious.
Interspecific hybrids are commonly seen in Weissia. Khanna(1960) opined that Weissia exserta (n =
26) is an allopolyploid derivative of W. cripta X W. controversa, both having n = 13. Pettet (1964) has
shown the possibility of interspecific as well as intergeneric hybrids in Funariaceae. Smith (1978) has
also shown the possibility of hybridization between Bryum pendulum and B. caespiticium. Molecular
studies of Ripetski (1992) also show that “level of interspecific genetic variability of mosses, is higher
than traditionally expected”.
338 � Bryophyta
CYTOTAXONOMY 29.9
The use of chromosome studies (i.e. number, structure and behaviour) in taxonomic work is known as
cytotaxonomy. In several mosses, the cytological studies have been used to determine relationships
at different levels confirming definite role of cytotaxonomy. A few selected examples of the use of
cytotaxonomy in bryophytes are listed below:
1. Anderson and Bryan (1956) separated closely related species of Fissidense cristatus (n = 12 +
1) and F. adianthioides on the basis of their cytology.
2. Yano (1957) discussed chromosome numbers and chromosome morphology of 22 families
including 63 genera and 139 species and made karyotypic comparisons of taxa. He opined that
members of Fissidentales, Grimmiales, Dicranales, Eubryales and Polytrichales have unique
karyotypic characters.
3. Mehra and Khanna (1961) worked on the cytology of mosses. They proposed that their
chromosome numbers support their classification based on the structure of peristome. The
mosses, possessing a single peristome, differ in range of chromosome number than those
possessing double peristome.
4. Kumar (1966), and Smith and Newton (1966) studied the chromosome morphology and
chromosome number of Amphidium lapponicum and Ptychomitrium polyphyllum and sorted
out their taxonomic misplacement by placing them in Grammiaceae instead of Isobryaceae.
5. Smith and Newton (1968) studied Polytrichales and observed that all counts in this order of
mosses are referable to n = 7, 14 and 21. They also opined that Tetrapidales have “evolved by
way of simplification from Polytrichales”.
6. Steere (1972), Newton (1972) and Ramsay (1974) worked on the cytotaxonomy of several
mosses. Chromosome number in Dicranales, Eucalyptales, Rottiales and Grimmiales are n
= 12, 13 and 14. Low chromosome number (n = 5 or 6) has been reported in members of
Fissidentales. In Tortula papillosa, the chromosome number is n = 7 (Newton, 1972) or n = 6
+ 1m (Ramsay, 1974).
7. Smith (1978) proposed a phyletic scheme of Bryopsida on the basis of the chromosome
numbers. He suggested that members of Polytrichales, Dicranales, Fissidentales, Funariales
and Orthotrichales show definite trends while those of Grimmiales, Pottiales, Isobryales,
Thuidiales, Hookeriales and Hypnobryales do not show any definite trend in the chromosome
numbers.
1. Give a precise account of cytogene cs of bryophytes in about 1000 words.

2. Feulgen staining technique is used for studying
(a) morphology, (b) cytology, (c) gene cs, (d) embryology.
3. Aceto-orcein stain gives good results for studying _____ materials of bryophytes.
4. Give a brief account of any one of the following in about 200 words:
(a) Some studies on mitosis in bryophytes,
(b) Some studies on meiosis in bryophytes
5. With the help of only one sentence, differen ate between the terms (a) heteropycno c, and
(b) heterochroma n.
6. What do you mean by ‘M-bivalent’ or ‘special bivalent’?
7. Write a detailed botanical note on heterochroma n in bryophytes in about 200 words.
8. Differen ate between (a) chroma n, and (b) heterochroma n in about 100 words.
9. What is euchroma n?
10. Give a detailed account of chromosome number in all the three classes of Bryophyta.
11. What are sex-associated chromosomes? Describe them briefly with special reference to
bryophytes.
12. The term ‘h-chromosomes’ refers to _____ chromosomes.
13. Sex-associated chromosomes are found in most of the _____ species of mosses.
14. What are nucleolar chromosomes and accessory chromosomes? Describe them in brief with
par cular reference to bryophytes.
15. Write an essay on polyploidy in bryophytes in about 500 words.
16. Describe hybridiza on in bryophytes in about 100 words.
17. A polyploid species with sets of chromosomes from two or more different species is called
_____, and this phenomenon is called ______.
18. Write a note on cytotaxonomy in bryophytes in about 200 words.
30
Ecology of
Bryophytes
VIEWS OF JOHANNES ET AL. 2007 ON
ECOLOGY OF BRYOPHYTES 30.1
Johannes et al. (2007) published an article in Annals of Botany (Vol. 99) on cryptogam ecology with
particular emphasis on bryophytes, and opined that ecology of bryophytes “has the potential to meet
some of the important challenges of understanding and predicting the biogeochemical and climate
consequences of large-scale environmental changes”. Relevant data for certain biogeochemistry-related
characters are available for bryophyte “species in the literature, including those involved in bryophytic
nutrition and nutrient recycling, their anti-herbivore defences, and their potentials for carbon gain
and losses”. However, it may be said that still “very little is known about the role and applicability
of functional traits” of bryophytes with respect to biogeochemical cycling. Yet, bryophytes are
“paramount determinants of biogeochemistry in several biomes, particularly cold biomes and tropical
rainforests, where they (i) contribute substantially to above-ground biomass; (ii) host nitrogen-fixing
bacteria providing major soil N input; (iii) control soil chemistry and nutrition through accumulation of
recalcitrant polyphenols and through their control over soil and vegetation hydrology and temperature;
(iv) promote erosion and also prevent it (biological crusts in deserts); (v) provide staple food to
arthropods, with important feedbacks to soil and biota; and (vi) facilitate and compete with vascular
plants” (Johannes et al, 2007).
LIMITS OF ECOLOGY OF BRYOPHYTES 30.2

Ecology is the study of organisms in relation to their environment. Ecology of bryophytes may be
studied in a number of directions, but here, only a few selected aspects are briefly discussed.
1. Autecology
2. Synecology
3. Succession
4. Factors of the habitat
Ecology of Bryophytes � 341
AUTECOLOGY 30.3
The ecology of a single species in a habitat is known as autecology. Tamm (1953) made a significant
contribution on the autecology of Hylocomium splendens, a moss. He studied the habitat, growth,
yield and mineral economy of this species of moss. H. splendens forms an almost pure ground cover
in coniferous forests of Scandinavia. Its growth depends on the canopy of the coniferous trees. Annual
yield of this moss decreases regularly with the distance from the canopy of the tree. Distance from the
tree canopy also shows its effect on the nutrient concentration in this species of moss.
Tallis (1958) studied the autecology of widely distributed species of a moss (Rhacomitrium
lanuginosum). This study shows a particular combination of quite large-scale environmental factors. In
the tropical regions, this moss is “restricted to rocks on the higher mountains but its lower altitudinal
limit drops progressively northwards until in Northern Oceanic regions, and in the Arctic, it may be
abundant at sea level”. Its typical growth form is a group of lateral branches arranged in definite zones
along the main axis, and according to Tallis (1959), “only one zone is formed each year”. Annual
growth rate varies from 5 to 15 mm and never exceeds 20 mm per annum in British conditions. Due to
its very slow growth in calcareous grasslands, it is usually absent in such habitats. It is so because it (R.
lanuginosum) is unable to compete with much faster-growing higher plants. Rainfall, air humidity and
temperature are the major factors which govern the geographical distribution of R. lanuginosum.
Autecological studies of bryophytes are of definite use from various points of view. Presence of
some larger bryophytes serve as guides to forest authorities regarding the conditions and characters
of land. Copper mosses are of particular interest in establishing the prospects of particular ores in
the region. Many bryophytic species act as indicators. Polytrichum and Rhacomitrium are invariably
acid indicators, whereas Detrichium flexicaule and Encalypta streptocarpa are indicators of base-rich
conditions.
With reference to bog ecology, Sphagnum is of outstanding importance. About a dozen species of
this moss play essential roles in bog ecology as elucidated by Ratcliff and Walker (1958).
SYNECOLOGY 30.4
The ecology of all the organisms found in a habitat or an ecosystem is known as synecology. Synecology
of bryophytes may be studied by (i) recognition of communities of bryophytes; (ii) recognition of
different growth forms among bryophytes; (iii) applying quantitative methods in describing vegetation
of bryophytes; (iv) studying succession process and the time required for its completion; and
(v) studying habitat conditions.
30.4.1 Recogni on of Communi es of Bryophytes

The group of species of plants, animals, or both, living in the same habitat and interacting with each
other is known as community. A bryophytic community includes several species of bryophytes. For
example, a mossy covering containing several species (e.g. Brachythecium ratabulum, Bryum capillare,
Ceratodon purpureus, Cryphaea heteromalla, Frullania dilatata and Hypnum cupressiforme) represents
a bryophytic community. All these species in a community (i) compete with each other, (ii) react upon
one another, (iii) possess different morphology and growth rate, and (iv) survive with each other as
342 � Bryophyta
efficiently as possible. The change that is observed or seen in a particular bryophytic community is a
very slow process. It may take decades for a particular bryophytic community in its various stages of
succession prior of attaining a climax.
For more details, the readers may consult Phytosociology and Ecology of Cryptogamic Bryophytes
by Barkman(1958).
30.4.2 Recogni on of Different Growth-Forms among Bryophytes

Bryophytic vegetation can also be analysed by studying growth-form representatives of a community.
Five categories of growth-forms among bryophytic community, recognised by Gimingham and
Robertson (1950) and Gimingham and Birse (1957), are cushions, turfs, canopy-formers, mats and
wefts.
1. Cushions include two types of cushions, namely large cushions (e.g. Leucobryum) and small
cushions (e.g. Grimmia).
2. Turfs include tall turfs (e.g. Polytrichum commune), short turfs (e.g. Bryum argenteum) and
open turfs (e.g. Polytrichum aloides).
3. Canopy-formers (e.g. Mnium undulatum).
4. Mats include four types, namely rough mats (e.g. Eurhynchium striatum), smooth mats (e.g.
Frullania tamarisci), thread-like mats (e.g. Eurhynchium praelongum) and thalloid mats
(e.g. members of Marchantiales and Metzgeriales).
5. Wefts including rhizoid-free weft (e.g. Hylocomium splendens) and wefts with frequent
tufts of rhizoids (e.g. Thuidium tamariscinum).
A sixth category of pendulous forms has been erected by Iwatsuki (1960), a Japanese bryologist.
It is represented by genera such as Barbella, Pseudobarbella and Floribundaria. A community in
different types of habitats, however, exhibits different growth forms. In a community occurring
near water, four successive zones may be seen as the distance from the water increases. These are
smooth-mat forms (e.g. Eurhynchium riparioides), replaced successively by rough-mat forms (e.g.
Cratoneura filicinum), thalloid-mat forms (e.g. Conocephalum conicum) and dendroid forms (e.g.
Thamnium alopecurum). Impact of soil type and groundwater level on the composition of bryophytic
communities has been studied by Gimingham and Brynard (1960).
30.4.3 Applying Quan ta ve Methods in Describing Vegeta on of Bryophytes
Several recent studies of applying quantitative methods in describing bryophytic vegetation are now
available in the literature, but a few pioneer ones are those of Hope-Simpson (1941), Jenny (1941),
Cornish (1954) and Perring (1959, 1960)
Hope-Simpson (1941) studied the vegetation of English chalk grassland with particular emphasis
on bryophytes and lichens and also compared their occurrence in other calcareous grasslands. His
studies published in the Journal of Ecology are focused on the bryophytic flora in an exact quantitative
form. These studies, however, give no indication of the size of the area surveyed and also show least
emphasis on the account of the habitat and height of the turf. Eight species considered as of outstanding
importance in chalk grassland are Acrocladium cuspidatum, Camptothecium lutescens, Dicranum
scoparium, Fissidens taxifolius, Hylocomium splendens, Pseudoscleropodium purum, Rhytidiadelphus
squarrosus and R. triquetrus.
Cornish (1954) studied the origin and structure of the grassland types of Central North Downs.
Her studies on bryophytic communities has been published in Journal of Ecology and are based on
physiography and past history of the bud.
Jenny (1941) proposed the concept of independent variable to study the chalk grassland of Britain,
and his studies have been supported and extended by Perring (1959, 1960). Jenny’s method of study
include “changes in soil and vegetation values in relation to changes in one independent variable whilst
keeping all the others constants”. For studying bryophytic vegetation, Perring gave due emphasis
to topography, i.e. sensitivity of particular species to different slopes. Perring (1960) also gave due
consideration to factors such as humidity, pH, carbonates, and exchangeable calcium, potassium and
phosphates.
30.4.4 Studying Succession Process and Time Required for its Comple on
The process of development of vegetation, involving changes of species and communities with time
is known as succession. It occurs because the growth of plants alters the biotic and edaphic factors
of a habitat, making possible the colonisation of other species, Doignon (1949) of France studied
succession of bryophytes on rotting logs, and concluded that so gradual and slow are the various stages
of succession that it may well be over 30 years before the same type of bryophytic flora is restored.
A clear picture of succession of bryophytes on a living tree was given by Barkman (1958). The
pioneer colonisers were lichens, and after about 25 years appeared the liverwort Frullania dilatata.
Within a few years now developed the species of Ulota, Orthotrichum and some more genera. Then after
about 10 years developed mosses like Neckera and Anomodon and liverworts like Porella platyphylla.
The mature tree was finally inhabited by bryophytes such as species of Leucodon and Zygodon after
about 40 years. In several cases, it takes 100 to 120 years for the completion of succession process of
bryophytic communities.
30.4.5 Studying Habitat Condi ons

The place or kind of place, in which an organism, community or association is found, is known as
habitat. Mosses and liverworts are important components of the flora on the trunks and branches of trees,
which thus function as their habitat. A detailed account of habitat of bryophytes may be gathered from
Phytosociology and Ecology of Cryptogamic Bryophytes of Barkman (1958). Bryophytic vegetation
on the branch or trunk of trees is affected by (i) daily movements of the sun and availability of sunlight,
(ii) direction of the prevailing wind in the region, and (iii) availability of the water supply or moisture.
Nutrient supply and acidity are also important for the bryophytic flora in a particular habitat. A rich
epiphytic flora is available on the trees having high nutrient status and neutral pH. A habitat with salty
water is generally avoided by mosses, and almost none of them lives in sea water. Funaria hygrometrica,
however, can grow where the concentration of salts is permanently very high. A halophytic community
of bryophytes consisting of mosses (e.g. Grimmia maritima, Hypnum cupressiforme, Pottia heimii and
Totella flavovirens) has been found to be salt-tolerant by Shacklette (1961). Shacklette’s findings have
been published in Vol. 64 of Bryologist.
SUCCESSION 30.5
As mentioned earlier under Article 30.4.4, succession is the process of vegetation development,
involving changes of species and communities with time. Succession proceeds towards the natural
climatic climax where some kind of stability is reached. Ecologists have confirmed that Funaria
344 � Bryophyta
hygrometrica is a pioneer cononiser, and it is followed in sequential stages of succession by Ceratodon

purpureus and then Polytrichum juniperinum and P. piliferum along with Cladonia, a lichen. The entire
process of succession takes several years to complete.
Secondary succession of terrestrial bryophytes in New Jersey, USA, has been studied and described
by Bard (1965). The species present in relative abundance in the initial stages were Physcomitrium
turbinatum, Pleuridium subulatum and Weissia controversa. A leafy liverwort hydrosere has been
described on the Yakobi Island, Alaska, by Shacklette (1965), and his study has been published in Vol.
46 of Ecology. Two liverworts of hydrophytic nature of the initial stages of the succession process were
Nardia compressa and Scapania paludosa, and the physiographic climax community was achieved by
Cladothamnus pyroliflorus. Quarterman (1949) observed that a “Frullania-Orthotrichum stage initiates
succession of bryophytes on the bark of cedar trees.”
FACTORS OF THE HABITAT 30.6

Only major climatic factors (light, temperature and water), and edaphic factors with reference to
bryophytes are briefly discussed.
30.6.1 Clima c Factors

1. Light Rocks and the moss communities growing on them are exposed to high light intensities.
Mosses growing in caves show tolerance to low light intensities. It is, however, very difficult to sepa-
rate the effects of light from those of temperature and humidity factors. Generally, it has been observed
that bryophytes are less shade-tolerant than many algae and more so than many higher plants. March-
antia polymorpha has a tendency to tolerate a very deep shade. An example of shade-tolerant moss is
Thamnium lemani. Few bryophytes are highly shade-tolerant and restrict themselves only in caves, e.g.
Cyathodium cavernarum and Schistostega osmundacea. Extreme light intensities have lethal effects on
such bryophytes. Many species of Sphagnum are intolerant of shade.
2. Temperature Much researches have not been done on the maximum temperature limit on which
bryophytes can survive, but a few mosses (e.g. Barbula revoluta and Tortula muralis), have been seen
to sustain a temperature as high as 52°C. Clausen (1964) studied the tolerance of liverworts to tempera-
ture and desiccation. Many Hepatics can survive for weeks together in a temperature as low as –10°C.
Some bryophytes have been seen to survive also in freezing environments as low as –40°C for more
than a day (e.g. Frullania tamarisci, Gymnomitrion corallioides and Ptilidium pulcherrinum).
3. Water Bryophytes are mostly terrestrial, found in moist surroundings, and only a few members
are found in the water (e.g. Riccia fluitans). Corticolous mosses absorb moisture mainly from the atmo-
sphere. Terrestrial members fulfill their water requirements from both soil and atmosphere. There exist
reports which suggest that Sphagnum can absorb moisture 16–25 times of its own dry weight when its
surrounding atmosphere is supersaturated. Bryophytes also adapt drought-resistant features, if water is
relatively scarce or evaporation is excessive. Some examples of drought-resistant bryophytes include
Lunularia cruciata, Torula ruraliformis and Anoectangium compactum. It is actually the humidity
which has a profound effect on the distribution of liverworts. Several species of bryophytes occur on
Indian hills because of abundant moisture.
30.6.2 Edaphic Factors

Edaphic factors include the effects of the soil in an ecosystem. Bryophytes grow commonly on a variety
of substrates including moist soil, humus, rocks, etc.
Along with moist soil, humus is a factor of primary importance. Technically, humus is the layer
of organic matter at the top of a soil profile. It is the habitat of most decomposers. Many mosses
growing on rocks grow actually on humus. Some bryophytes prefer to grow on a particular kind of
humus, e.g. Dicranella cerviculata shows preference to acid peat, and Tetraphis pellucida to decayed
woods. Humus-loving mosses are generally saprophytic, and it is believed that microorganisms found
in humus are an important factor for these bryophytes.
Bryophytes, which are able to grow on calcareous or other mineral-rich substrata, belong to particular
families such as Tortulaceae and Amblystegiaceae. They also serve as indicators for the presence of
particular elements, e.g. copper. Nitrogen substances in the soil are essential for growth of bryophytes.
Certain mosses (e.g. members of Splachnaceae) are obligately nitrophilous. Some other nitrophilous
mosses are Lunularia cruciata and Bryum argenteum.
For the growth of many bryophytes, the hydrogen-ion concentration of the substratum is also an
important factors. Lime has been determined as the commonest cause of hydrogen-ion concentration
for bryophytes in the soil. Different species of Sphagnum were grown in water-culture solutions
maintained at different hydrogen-ion concentrations by Olsen (1923).
Mosses generally avoid salty water. But a few mosses are also halophytes and grow easily in salt-
tolerant places, e.g. Grimmia maritima and Tortella nitida. Epiphytic mosses generally obtain their
nitrogen from the humus of the bark on which they are developing.
30.6.3 Bio c Factors

The effects of living organisms on an ecosystem and on each other are studied under biotic factors.
Much studies have not been done on this aspect of ecology of bryophytes. Since moss carpets have the
power to retain water, they play an important role for seed beds of flowering plants on rocks. Mosses
also function as pioneers on sand dunes where they are replaced soon by lichens. Bryophytes are
always at a great disadvantage in competition with vascular plants mainly because of the small size
and restricted growth of the former. Bryophytes get suppressed easily by the dead remains of higher
vegetation.
1. Write an essay on ecology of bryophytes in about 1000 words.

2. Ecology of single species in a habitat is known as _____.
3. What is synecology? Describe briefly the synecology of bryophytes in about 500 words.
4. Define the term ‘community’ with reference to ecology of bryophytes.
5. What do you mean by succession? Describe succession process with reference to ecology of
bryophytes.
6. The place, in which an organism, community or associa on is found, is known as _____.
7. Give an account of clima c and edaphic factors with reference to the ecology of bryophytes.
31
Origin of
Bryophytes and
Their Fossil History
ORIGIN OF BRYOPHYTES 31.1
Bryophytes are nonvascular plants, mainly terrestrial in habitat, generally susceptible to desiccation,
and show heteromorphic alternation of generations, with dominant haploid gametophytic generation
and ephemeral sporophytic generation. Our knowledge of the geological history of this group of plants
is fragmentary because much is still not known about their fossil records. However, Walton (1928)
has shown the presence of bryophytes in the Upper Carboniferous, i.e. about 285,000,000 years ago.
Theories regarding the origin of bryophytes are, therefore, mainly based on our studies on ontogeny,
comparative morphology, and also analogies of bryophytes with other groups of living land plants.
Two main theories about the origin of bryophytes exist. Of these, one suggests that they are
descendents of pteridophytes while the other suggests that bryophytes have their origin from algae.
Before discussing some details of both these theories, let us first examine a few details about the
monophyletic or polyphyletic nature of this group.
A NOTE ON THE MONOPHYLETIC OR POLYPHYLETIC

NATURE OF BRYOPHYTES 31.2
Based on morphological characteristics, workers like Campbell (1895) and cavers (1911) suggested
bryophytes to be a monophyletic group. Even phylogenetic trees were constructed on the basis of these
characteristics. Satisfactory details of cytology were, however, not available to bryologists in those
days, as are available today, and hence the picture was not clear. The theory suggesting bryophytes to be
a monophyletic group is, therefore, now nothing more than history. It has been rejected by later workers
including Schuster (1966), Chopra (1967) and Crum (1976). A majority of the later workers now believe
in polyphyletic origin of bryophytes. But, Fulford (1964) remarked that since detailed information is
still not available on a majority of bryophytic genera, all these suggestions about monophyletic or
polyphyletic nature of bryophytes are uncertain and tentative, and much more is still to be done in this
regard by bryologists.
Origin of Bryophytes and Their Fossil History � 347
Geiger (1990) and others recognised four distinct lines within bryophyta, namely Hepaticopsida,
Anthocerotopsida, Bryopsida and Sphagnopsida. All these are individual natural groups with clear
differences in their morphology, anatomy, ontogeny of their sex organs, cytology and chemical
structure, e.g. biflavonoids are of frequent occurrence in the cell walls of Bryopsida and Sphagnopsida
while they have not been isolated from Hepaticopsida and Anthocerotopsida. Michler et al. (1992)
employed molecular data to suggest polyphyletic origin of bryophytes. They analysed chloroplast-
coded 16S and 23S-rRNA genes of several algae and bryophytes including liverworts, hornworts and
mosses, and on these basis, they supported the polyplyletic origin of bryophytes.
Further details of phylogeny are discussed in a separate chapter.
ORIGIN OF BRYOPHYTES FROM PTERIDOPHYTES 31.3

Discovery of Psilophytales by Kidston and Lang (1917–1921) suggested that bryophytes are descendants
of pteridophytes. Psilophytales are some of the simplest and probably the oldest pteridophytes, not
existing today. The sporophyte of Psilophytales (e.g. Rhynia, Horneophyton) was a rootless, leafless,
dichotomously branched shoot bearing terminal sporangia. These pteridophytes could be compared on
one hand with bryophytes (e.g. Anthocerotales) and, on the other hand, with vascular plants. However,
there existed several differences between bryophytes and Psilophytales, such as (i) the sporophyte was
physiologically independent in Psilophytales and not in bryophytes, (ii) continued apical growth
was shown by Psilophytales and not by bryophytes, and (iii) vascular tissue (xylem and phloem) was
present in Psilophytales and absent in bryophytes. There also exist several similarities between
Psilophytales and bryophytes. The sporangia of some Psilophytales (e.g. Rhynia, Horneophyton)
resemble the capsules on several bryophytes (e.g. Sphagnum, Andreaea, and some Anthocerotales).
Proskauer (1960) has shown several similarities between capsules of Anthocerotales and Rhyniaceae
of Psilophytales. He observed remarkable similarity between the sporangia of Horneophyton
(of Psilophytales of Pteridophyta) and thickened columnar surface layer and jacket lining layer of
Dendroceros crispus (of Anthocerotales of Bryophyta). Presence of tracheid-like elements with
spiral thickenings in columella of Anthoceros also support the pteridophytic origin. Due to such close
similarities, botanists opined that bryophytes (especially Anthocerotopsida) may have originated from
simplest known vascular plants, such as Psilophytales of pteridophytes.
Two main advocates of pteridophyte ancestry of bryophytes are Haskell (1949) and Christensen
(1954). Haskell (1949) opined that bryophytes “represent a group originating from Psilophytalean
ancestry, following reduction due to their habitat”. Haskell’s researches have been published in Vol.
12 of the journal Bryologist. Christensen (1954) published his detailed observations in Vol. 51 of Bot.
Tidsskr where he commented that “bryophytes have descended from pteridophytes”. Proskauer (1960),
in his series of publications in Phytomorphology, suggested that “Anthocerotopsida originated from
forms such as Horneophyton” of pteridophytes. The Indian bryologist Shiv Ram Kashyap (1919) also
believed that due to several common features, “Hepaticopsida may have arisen from pteridophytes.”
Pteridophytic ancestry of bryophytes was also postulated by Kashyap and Dutt (1925). Jennings (1928)
and Schwarz (1955) have also favoured the idea of bryophyte-origin by way of simplification from
Psilophytales.
Lignin is the chemical constituent of tracheids. It is absent in bryophytes. However, its occurrence
in the gametophytes and peristome of some mosses (e.g. Dawsonia) suggests their origin from
348 � Bryophyta
pteridophytes (Siegel, 1962, 1969). Niklas and Pratt (1980), however, have questioned the presence of
lignin in bryophytes.
ORIGIN OF BRYOPHYTES FROM ALGAE 31.4

31.4.1 View of Lignier (1903)
Living bryophytes show amphibious nature and due to this, it was suggested by earlier bryologists that
bryophytes have originated from aquatic ancestors, like algae. Lignier (1903) was the first to suggest
that algal ancestors of bryophytes gave rise to a primitive terrestrial type, the Prehepatics, which gave
rise to bryophyta, on one hand, and vascular cryptogams, on the other.
31.4.2 View of Bower (1908)

Archegoniatae are plants having archegonia, e.g. bryophytes, pteridophytes and gymnosperms. In
his theory of origin of land flora, Bower (1908) proposed that Archegoniatae originated from aquatic
ancestors (e.g. algae) “inhabiting shallow freshwater or higher levels between marine tide-marks”.
Amongst the inhabitants of aquatic habitat, bryophytes show resemblances with Chlorophyceae (green
algae) due to several features, including (i) the presence of photosynthetic pigments in the assimilatory
tissue, (ii) the presence of starch as the metabolic product, and (iii) the presence of cell wall made up
of cellulose. Thus, it was concluded that Chlorophyceae were probably the nearest algal relative of
Bryophyta.
31.4.3 View of Fritsch (1935, 1945)

F E Fritsch (1935, 1945), a well-known algologist, believed that members of the order Chaetophorales
of the class Chlorophyceae contain all characters for the development of more complex types, and
“evolution of the archegoniate plants from algae may have followed” the undermentioned course of
actions:
1. Development of heterotrichous habit showing formation of different types of filaments or
trichomes “in the plane of substratum, i.e. prostrate and in the upright vertical direction,
perpendicular to the prostrate system” as in Chaetophorales of Chlorophyceae.
2. Development and formation of parenchymatous structures in the upright filaments as in algal
members of Fucales and Laminariales.
3. Initiation and ultimate establishment of apical growing point in the upright filaments, as in
Chaetophorales, Fucales, Laminariales, etc.
4. Disappearance of prostrate system and establishment of dichotomous branching in the algal
members.
5. Establishment of vascular system in the upright aerial parts, e.g. scalariform thickenings of
medullary cells of Sargassum and sieve-tube-like elements in Macrocystis.
6. Development of cuticle outside the epidermal layer.
Development of protonema in the mosses (Bryopsida) was seen to be equivalent to the prostrate
system and of leafy gametophores to the erect filaments of Chaetophorales (algae) by Fritsch (1945).
Fully developed heterotrichous habit has been later observed in the developed protonema of mosses.
31.4.4 Some Other Views on the Algal Origin of Bryophytes

The origin of bryophytes from algae, particularly green algae, has also been suggested and supported
by workers like Schuster (1966), Chopra (1967), Fott (1974) and Karunen ((1990). It is mainly due to
the resemblances in their (i) photosynthetic pigments, (ii) cell-wall components, (iii) flagella, and (iv)
food reserves. According to Watson (1971), recapitulation of ancestral algal habit can be seen in the
filamentous protonema of mosses. Many biological, physiological, reproductive and developmental
features also suggest the green-algal ancestry of not only of bryophytes but also of other land plants. It
is also evidenced by fossil records that some present-day green algae (e.g. Coleochaete, Chara, etc.) are
quite ancient, extending back to Silurian, i.e. approximately 400 million years ago.
Amongst green algae, Charales (e.g. Chara) are generally linked with ancestry of land plants. This
is due to the presence of several specific features that are common to both. Only advanced Charales
among algae possess phragmoplast, which are also characteristic of vascular plants and bryophytes.
(Phragmoplast is a complex of interdigitating microtubules aligned more or less parallel to the
earlier spindle microtubules, developing at the end of mitosis). A majority of other green algae lack
phragmoplast. Brown and Lemmon (1990) have also advocated the algal origin of bryophytes. Graham
(1980) also opined that Charales also show similarity with other land plants (including bryophytes,
pteridophytes and gymnosperms) in microtubular cytoskeleton of their flagellum. Photorespiration in
Charales is also similar to the photorespiration in land plants.
Phytochrome is a pigment which controls many of the physiological responses of plants to light. It
absorbs the red and far-red wavelengths of light. Scientists have proved the similarity of phytochrome
between Charales and land plants.
Hori et al. (1985), on the basis of their molecular studies, opined that “Nitella-like green plants may
be the direct ancestors of first land plants”, in general, and bryophytes in particular.
FOSSIL HISTORY OF BRYOPHYTES 31.5

Due to the fragile and delicate nature of the plant body of bryophytes, we have a very limited knowledge
of their fossils. These plants also do not possess a cutinised epidermis and lignified vascular tissues
like xylem and phloem. In spite of this, there are some authentic records of fossil liverworts in Upper
Devonian and mosses in Carboniferous. Hepatic characters have also been noted in earliest land plant
fossils from Ordovician and Silurian, i.e. much before the advent of the vascular plants, by Taylor (1995)
and Edwards et al. (1995). Parihar (1987) mentioned that “the fossil Bryophyta do not throw any light
on the origin and evolution of the group”. A preliminary account of some known fossil Hepaticopsida
and Bryopsida is given here.
31.5.1 Fossil Hepa copsida
1. Hepa cites Walton (1925) was the first to provide authentic details of the relationship of some
fossil plants of Upper Carboniferous with some living Hepaticopsida, and these have all been assigned
to the form-genus Hepaticites, a leafy liverwort (Fig. 31.1A). No reproductive structure of Hepatic-
ites was reported by Walton (1925), who compared the species of this form-genus with anacrogy-
nous Jungermanniales of Hepaticopsida. Of the five species of Hepaticites described by Walton (1925,
350 � Bryophyta
1928), H. kidstoni (Fig. 31.1A) had a “broad axial region and two definitely arranged series of leaves
or lobes with two accompanying series of smaller scale-like appendages”. Due to these characters, H.
kidstoni resembled the living genus Treubia. The other two species (H. langi and H. willsii) resembled
with the living genus Riccardia due to their parenchymatous thalli with no midrib. The remaining two
species described by Walton are H. lobatus and H. metzgerioides. The former had an axial region and
the lobed wing, while the latter resembled closely the living genus Metzgeria, hence named Hepatites
metzgerioides. Heuber (1961) described H. devonicus from Upper Devonian. It had a rhizome-like
portion bearing unicellular rhizoids and a dichotomously branched thalloid portion bearing wings and
thick midrib. Schuster (1966) shown resemblances of Hepaticites devonicus with the present-day Pal-
lavicinia and renamed it as Pallavicinites devonicus.
Fig. 31.1 Some fossil bryophytes: A, Hepa cites kidstoni; B, Leafy shoot of Naiadita;
C, Sec on of sporogonium of N. lanceolata showing foot, capsule and spores
(A, a er Walton; B-C, a er Harris)
Harris (1931) described three species of Hepaticites (viz. H. glebas, H. laevis and H. rosenkrantzi)
from Jurassic of Greenland, and later on four species (H. arcuatus, H. haiburensis, H. hymenoptera and
H. wonnacotti) from Jurassic of Yorkshire (Harris, 1961). Two more species described by Harris are H.
amauros and H. solenotus from Triassic of Greenland.
2. Naiadita Naiadita (Fig. 31.1B) is the most completely known fossil representative of bryophytes
reported from Upper Triassic of England by Harris (1938), who opined that it is quite close to Riellace-
ae of Sphaerocarpales. Harris described almost all parts (rhizoids, stem, leaf, gemma cups, archegonia,
embryo, sporophyte and spore) of this fossil bryophyte, except antheridia. Sparsely branched stem of
N. lanceolata was 1–3 cm in height. Basal part of the stem had several unicellular rhizoids. A major
part of the stem had parenchymatous cells, and there was no differentiation of tissues. The leaves on
the stem were linear in the lower portion while lanceolate to almost round towards the apical portion.
Gemma cups having multicellular gemmae were also present. Mature archegonia were surrounded by
leaflike lobes of perianth. The sporophyte (Fig. 31.1C) had a spherical capsule and small hemispherical
foot with no seta. Columella and elaters were absent while there were present many lenticular spores
in the capsule. Naiadita is unique in combining features of both liverworts and mosses. It also shows
some characteristics resembling that of Marchantiales and Calobryales.
3. Some Other Fossil Hepa copsida Lundblad (1954) described Marchantites hallei from
South America and Marchantiolites porosus, Ricciopsis florinii and R. scanica from the coal mines
of Skromberga (Sweden). Species of Ricciopsis are the forms like Riccia. Berry (1919) has earlier
described from Upper Cretaceous the forms having affinities with Jungermanniales acrogynae and
assigned them to belong to Jungermannites cretaceous. Townrow (1959) described Hepaticites cy-
athodoides from the Middle Triassic of Natal. It had several resemblances with Cyathodium.
31.5.2 Fossil Bryopsida

Some of the well-preserved fossil mosses belong to the species of Muscites, Intia and Protosphagnum.
Muscites bertrandi and M. polytrichaceous have been described from the Upper Carboniferous,
in the form of either compression of leafy shoots or small petrified shoots possessing rhizoids with
oblique septa. M. polytrichaceous had stems with small univeined leaves, which can be compared with
the present-day Polytrichum and Rhizogonium (Dixon, 1927).
Neuberg (1956, 1958, 1960) described well-preserved fossil mosses from Lower and Upper Permian
rocks of Angaraland of former USSR. Intia vermicularis (Fig. 31.2A) and I. variabilis (Fig. 31.2B) are
two such mosses, remarkably similar to present-day Bryum and Mnium. Other species of fossil mosses
described by Neuberg are Intia falciformis, I. angustifolia, Salairia longifolia, Uskatia conferta,
Polyssaievia spinulifolia, Buchtia ovata and Bajdaievia linearis.
Fig. 31.2 Some fossil mosses A, In a vermicularis showing a leaf and part of stem;
B, I. variabilis showing part of the leaf enlarged; C, Part of leaf of Protosphagnum
nervatum near nerve and leaf cells (A-C, a er Neuberg)
Fossils of the leaf genus Protosphagnum (Fig. 31.2C) are similar to present-day Sphagnum. They
possessed a network of two types of cells arranged in a specific manner, in which dividing colourless
cells are clearly visible. A new order Protosphagnales has been created by Neuberg (1960). Three
352 � Bryophyta
genera included in this order are Jungagia, Protosphagnum and Vorcutannularia. These genera possess
two kinds of cells in their leaves, thus resembling closely the present-day Sphagnum. Gam (1962)
criticised the inclusion of these three genera under an independent order Protosphagnales and has tried
to establish their affinity with orders like Polytrichales and Dicranales.
Fossil mosses were of rare occurrence in the Mesozoic. Berry (1928), however, reported Muscites
lesquereuxi from the Late Cretaceous of North America, and Townrow (1959) described M. guescelini
from Triassic of Natal.
Many members of Bryopsida, reported from Cenozoic include the form-genera Polytrichites,
Plagiopodopsis and Palaeohypnum.
1. Write a detailed note on the monophyle c or polyphyle c nature of bryophytes in about 200
words.
2. Give an account of the origin of bryophytes.
3. Bryophytes are descendants of pteridophytes. Elaborate this statement in about 250 words.
4. Explain various views about the origin of bryophytes from pteridophytes.
5. Discovery of which fossil order suggested that bryophytes are descendants of
pteridophytes?
6. Write an essay on the origin of bryophytes from algae.
7. Define the term ‘archegoniatae’.
8. Archegoniatae includes which of the following?
(a) Bryophytes, (b) Pteridophytes, (c) Gymnosperms.
9. Explain the view of F E Fritsch regarding the origin of bryophytes from algae.
10. Explain the following terms: (a) Phytochrome, (b) Phragmoplast
11. Give an account of fossil history of bryophytes in about 500 words.
12. Write short notes on:
(a) Hepa cites, (b) Naiadita.
13. Give a brief descrip on of some fossil Hepa copsida.
14. What do you know about fossil mosses? Explain them in some detail.
15. What is common amongst the following?
(a) In a, (b) Muscites, (c) Protosphagnum
These are _____ _____.
32
Economic
Importance of
Bryophytes
UTILITY OF BRYOPHYTES: A GENERAL OVERVIEW 32.1
The economic importance of bryophytes (liverworts, hornworts and mosses) is relatively unknown
to most people. But in different parts of the world, bryophytes are now widely used because they (i)
modify their microclimate, (ii) serve to conserve moisture, (iii) check soil erosion on hilly slopes, (iv)
serve as a seed bed for forest cover, (v) help in pollution monitoring, and also (vi) are now considered
as new sources of pharmaceutical products. Viewed from a broad perspective, bryophytes are used (i) in
horticulture, (ii) for household purposes, (iii) for their ecological importance, (iv) in preparing several
pharmaceutical products, and also (v) in the biomapping of atmospheric precipitation.
Some of the specific uses of these nonvascular plants are briefly discussed here in this chapter.
ECOLOGICAL USES OF BRYOPHYTES 32.2

Bryophytes have been found to be good indicators of environmental conditions, and this aspect of these
plants has been discussed in detail in Chapter 23.
MEDICINAL USES OF BRYOPHYTES 32.3

32.3.1 Some General Examples of Medical Uses
1. In the ancient times, various mosses (e.g. Bryum, Mnium, Philonotis) were crushed into a kind
of paste and applied as a poultice.
2. Burned ash of mosses, mixed with honey and fat, is used as an ointment for cuts and wounds
in several parts of our country.
3. Marchantia polymorpha and M. palmata are used as medicines for boils and abscesses by
Himalayan Indians.
4. A paste made from various species of Riccia is applied externally to cure ringworm.
354 � Bryophyta
5. Traditional medicines are made in China from over three dozen bryophytes to treat various
illnesses like bronchitis, tonsilitis, tympanitis, cardiovascular diseases and skin diseases.
6. Extracts made from Rhodobryum giganteum and R. roseum can enhance flow of blood in the
aorta by as much as 25% to 30% in some animals. These species are used to treat cardiovascular
diseases and nervous prostration.
7. A blood-clotting factor IX is produced from transgenic Physcomitrella and used in the
treatment of haemophilia-B.
8. Sphagnum has been used for various medicinal purposes. Calcified peat is very effective as
a germicide. Peat water has antiseptic and astringent properties. Sphagnol is a distillate of
peat tar and highly valuable in treating eczema, psoriasis, hemorrhoids, scabies, insect bites,
acne and other skin diseases. Dried Sphagnum is sold in China to treat hemorrhages (Bland,
1971).
9. Due to the presence of several biologically active compounds, bryophytes survive in nature,
in spite of the absence of thick cuticle and bark. These compounds protect bryophytes from
fungi and other microorganisms, and due to this, they are of medicinal value to humankind.
For details of biologically active compounds in bryophytes, refer Chapter 24.
10. Extracts of liverworts, such as Pallavicinia and Reboulia, are well-known for their antifungal
and antimicrobial activities. It is due to the presence of lunularic acid in these genera.
11. Antibiotically active substances have also been extracted from some mosses, such as
Polytrichum, Sphagnum and Atrichum.
12. Petroleum-ether extracts of Barbula and Timmiella are highly effective against Gram-negative
and Gram-positive bacteria.
13. Skin infections caused by bacteria such as Bacillus subtilis, and fungi such as Candida
albicans and Trichophyton mentagrophyte can be checked by the extract of Plagiochila
stevensoniana.
14. Extracts of some liverworts act as stomach poison in some animal pests (e.g. Arion
lusitanicus).
15. Certain compounds effective against leukaemia have been isolated from Plagiochila fasciculata.
Diplophyllin, a compound isolated from some species of the liverwort Diplophyllum, is
significantly effective against human epidermoid carcinoma.
16. Sphagnum is extensively used for dressing wounds. Its pads are preferred in place of cotton,
because they easily absorb liquids as much as four times than cotton, and are cooler, softer and
less irritating to skin than cotton.
17. Marchantia polymorpha was widely used in earlier times in China and India to treat liver
ailments. According to Bland (1971), it “cools and cleanses the liver, removes yellow jaundice,
and also removes inflammation”. It is also used to treat pulmonary tuberculosis in some parts
of Europe, according to Bland (1987). According to Hu(1987), it is still used in China “to treat
jaundice of hepatitis and as an external salve to reduce inflammation.”
18. Polytrichum commune is used to reduce inflammation and fever, as a detergent diuretic,
laxative and hemostatic agent according to Hu (1987). Plants of this species are boiled to make
a “tea” to treat common cold. It also dissolves stones of the kidney and gall bladder (Gulabani,
1974).
19. Haplocladium microphyllum is used to treat bronchitis, cystitis, tonsilitis and tympanitis.
Economic Importance of Bryophytes � 355
32.3.2 An bio cs from Bryophytes

Liverworts and mosses rarely show signs of infection in nature and it is mainly due to their characteristic
of “inhibition of microorganisms in products of bryophytes”, e.g. Sphagnum portoricense, S. strictum,
Conocephalum conicum and Dumortiera hirsuta (Madsen and Pates, 1952). McCleary et al. (1960)
opined that mosses are a definite source of antibiotics. Antimicrobial activity of the extracts of Porella,
Pallavicinia and Reboulia has also been confirmed by workers like Belcik and Wiegner (1980) and
Isoe (1983). Antibiotic properties in over a dozen mosses (e.g. Atrichum, Polytrichum and Sphagnum)
has also been investigatd by McCleary and Walkington (1966).They suggested that these mosses
“strongly inhibited either or both Gram-positive and Gram-negative bacteria”. Dicranum scoparium
strongly inhibited all bacteria except Escherichia coli. Species of Barbula and Timmiella also show
high occurrence of antibacterial activity according to researches of K G Gupta and B Singh (1971).
Ichikawa et al. (1983) tested nearly 80 species of mosses and “found antimicrobial activity in nearly
all of them”.
“Peat humic acids possess antiviral activity against herpes simplex virus types 1 and 2” (Klocking
et al., 1976). Extracts of Sphagnum and Camptothecium possess antiviral active humic acids that can
inhibit the growth of polio virus, according to Witthauer et al. (1976).
32.3.3 An -cancerous Proper es of Bryophytes

1. Extract of Polytrichum juniperinum has anti-cancer activity against Sarcoma 37 in mice
(Belkin et al., 1952).
2. A compound diplophylline, obtained from Diplophyllum albicans and D. taxifolium shows
anti-cancerous activity against human epidermoid carcinoma (Ohta et al., 1977).
3. Compounds, such as sesquiterpenoids, costunolide and tulipinolide, effective against
carcinoma of vasopharynx, have been isolated from Conocephalum supradecompositum,
Frullania monocera, Lepidozia vitrea, Marchantia polymorpha, Plagiochila semidecurrens
and Porella japonica by Asakawa (1982) and Matsuo et al. (1981, 1984).
4. Spjut et al. (1986) of the US National Cancer Institute tested over 200 species of bryophytes
(184 species of mosses and 23 species of liverworts) and found that extracts of 43 species
possess anti-cancerous properties. Most active of these bryophytes belong to Brachytheciaceae,
Dicranaceae, Grimmiaceae, Hypnaceae, Mniaceae, Neckeraceae, Polytrichaceae and
Thuidiaceae.
5. Anti-leukemic activity of Marchantin-A from Marchantia polymorpha and riccardin from
Riccardia multifida has also been reported by Asakawa et al. (1982).
BRYOPHYTES AS FOOD SOURCE 32.4

Bryophytes as food source for animals are not very important. However, “Alaskan reindeer occasionally
graze on Aulacomnium turgidum, Hylocomium splendens and Polytrichum” (Band, 1971). During
scarcity of fresh food, bryophytes may be the source of specific needs of some animals, e.g. Barbella
pendula has a high content of Vitamin B12, which is not easily available in many vegetarian diets. For
giving good vitamin and iron diet, piglets and some other animals are given a diet of milled peat moss
(Sphagnum). According to JH Bland (1971), the “Chinese consider mosses to be a famine food”. In
356 � Bryophyta
some European countries, Sphagnum contributes to the flavour of Scotch Whisky, according to N G
Miller (1981). According to G B Pant and S D Tewari (1989), Kumauni Indians use some bryophytes
(e.g. Anomodon, Entodon, Hypnum, etc.), wrapped in a cone of Rhododendron companulatum leaves,
to serve as a filter for smoking tobacco. Some aphids depend on mosses as food for their larvae. Since
mosses have very low calorific values, some scientists are now exploring this aspect of bryophytes.
HORTICULTURAL USES OF BRYOPHYTES 32.5

Since long, bryophytes have been used in horticulture as “soil additives, ground cover, dwarf plants,
greenhouse crops, potted ornamental plants and for seedling beds” (Sjors, 1980). Besides its common
use in making poles to support climbing plants, Sphagnum is also used for other decorative horticultural
purposes including “making baskets and covering flower pots and containers for floral arrangements”
(Thomason, 1994). Wet plants of Sphagnum are used in nurseries for shipping living plants. Gardeners
use Sphagnum in air layering, a method of propagating plants. In Japan, preparation of landscape types
is an alternative horticultural art, in which several mosses (e.g. Bartramia pomiformis, Leucobryum
neilgherrense and Polytrichum commune) are used. Mosses are also used as an important element for
bonsai, where they help to stabilise the soil and retain moisture.
HOUSEHOLD USES OF BRYOPHYTES 32.6

Bryophytes, especially mosses, are used for decorative purposes in many countries including Finland,
England, France, Japan and USA. Because of the absorbent and insulating properties, Sphagnum is
the most useful household moss. Dried plants of Climacium japonicum are used in Japan for making
ornamental white flowers. Mats are prepared and sold in the market in many parts of India. Beddings,
mattresses, cushions and pillows are prepared by stuffing mosses in many parts of India. In some
parts of Himalayas, mosses are used as insecticides and insect repellents while storing grains. A cheap
kind of cloth is prepared by mixing Sphagnum with wool in Germany. Head cushions for carrying vessels
of water and other heavy articles are prepared by several mosses including Hylocomium, Hypnum and
Trachypodopsis. Several bryophytes (e.g. Hypnum, Macrothamnium, Neckera, Sphagnum, etc.) are
used for packing of apples, plums and other fruits in India.
Climacium americanum is used or making wreaths and crosses. Moss roses are prepared from
Hylocomium splendens. Bryum is collected and used for floral arrangements in USA. Rhytidiadelphus
is used in craft display in hotels. Green carpets for floral exhibitions are prepared from species of
Hylocomium and Rhytidiadelphus. Several bryophytes are graceful looking than many aquatic higher
plants, and are, therefore, used in aquaria, e.g. Bryum pseudotriquetrum, Riccia fluitans, Ricciocarpus
natans, Fontinalis antipyretica and Rhacopilum aristatum (Takaki et al, 1982).
HOUSE CONSTRUCTION 32.7

Bryophytes are used in the construction of houses, furnishing, boats, and other items in the parts of the
world where woody plants are scarce and bryophytes are common. Moss mats are used with grasses,
shrubs and bamboo to make doors at the openings of huts in several parts of Himalayas. Peatcrete,
and peatwood, the products made from Sphagnum peat, are used as construction materials. Peat-based
pasteboard and wrapping paper is manufactured in the USA. In Phillippines, bryophytes are used as
fillers between wooden posts of walls and shingles of roofs (Tan, 2003). Fontinalis antipyretica has
been used as fire insulation between the chimney and walls in countries like Norway, Denmark and
Finland (Thieret, 1956). According to Pant and Tewari (1989), shepherds in Himalayan highlands
use several bryophytes as chinking in temporary summer homes. Some such bryophytes are species
of Actinothuidium, Anomodon, Entodon, Herbertus, Floribundaria, Philonotis, Plagiochila and
Trachypodopsis. In some countries of Europe, Sphagnum is stuffed between timbers of houses to deaden
sound. In Russia, pressed and heated slabs of Sphagnum are used to insulate houses and refrigerators
(Rue et al. 1977). Mosses are now being widely used throughout Germany as a roofing material.
Polytrichum commune is used to make nautical ropes in Belgium. According to Rue et al. (1977),
“peatcork is made from the coarse fraction of peat, while peatfoam is an ultra-light construction
material based on peatmoss and foamed resin”.
MOSS INDUSTRY 32.8

Being highly absorbent and permeable, Sphagnum can absorb metals. Due to these characteristics, it
is used as an effective filtering and adsorption agent for treating wastewater and effluents of factories
containing toxic discharge of heavy metals, e.g. silver, copper, cadmium, mercury, iron, lead, etc. It
is also used in filtering oils, dyes, detergents and even microorganisms. Active carbon, used in many
chemical industries, can also be produced from Sphagnum. Peatmoss (Sphagnum) is also used in
producing several high-quality products in the industries related to pollution control and bioremediation.
Several mosses have oil-absorbing qualities, and are therefore useful for oil spills on waterways. Some
bryophyte-derived products are used to absorb fluids containing acetone, benzene, chloroform, corn oil,
diesel fuels, gasoline, jet fuels, kerosin, oil-based ink, paraffin oils, xylenes, etc. Sphagnum is highly
useful because it has the ability to hold up to 30 times its weight in water. It is, therefore, an excellent
material for shipment of plants, flowers and fresh vegetables. It is used on a large scale for storage of
roots and bulbs, and also for hydroponics gardening. Insulator sheets for houses are also manufactured
from Sphagnum and other mosses. In France, moss carpets of various sizes are also prepared. These can
be easily fixed along lawns, roads, playgrounds, etc.
FUEL FROM BRYOPHYTES 32.9

In several European countries, including Finland, Sweden, Poland and West Germany, liverworts and
mosses are used as a fuel. Products like low and intermediate BTU gas, ethylene, hydrogen, methanol
and natural gas are produced from peat. Methane is produced on a large scale from peat mosses (e.g.
Sphagnum). The heating value of peat is said to be superior than that of wood. According to the reports
of United Nations (1981), “nearly half the world’s annual peat production is used for fuel, with peat
resources worldwide estimated to be equivalent to 100–200 million tons of oil, or about half the
known gas reserves”. Peat is said to be a clean-burning fuel. In the past few decades, peat has received
unprecedented attention as an alternative fuel source.
358 � Bryophyta
According to Boffey (1975), the “former Soviet Union burned an estimated 70 million tons, and
Ireland 3.5 million tons of mosses in one year to produce electricity”. Richardson (1981) opined
that about “25% of the fuel in Ireland is moss-based”. However, improved methods are needed for
harvesting, drying and conversion of peat mosses to a burning fuel, according to Lindstrom (1980).
Besides the above-mentioned uses of peat, “Finland is exporting pulverised peat to northern Sweden,
where use in industry and municipal heating, power generation, and oil burners of pulp and paper
companies is increasing” (Summerton, 1981).
PESTICIDES FROM BRYOPHYTES 32.10

According to Yepsen (1984), bryophytes contain natural pesticides in their body. Sesquiterpene
hemiacetyl plagiochiline, a poison extremely potent in mice, is present in the liverwort Plagiochila
according to Matsuo (1983). This pesticide also inhibits the feeding of an African army worm, according
to Asakawa et al. (1980). Pesticides, like ferulic acid and coumaric acid, have also been isolated from
Brachythecium rutabulum and Mnium hornum (Davidson et al, 1984). Asakawa (1990, 2001) also
isolated many terpenes and other phenolic compounds having pesticidal properties from species of
Porella.
CLOTHING FROM BRYOPHYTES 32.11

According to Hedenas (1991), Sphagnum is used in Germany “to line hiking boots, where it absorbs
moisture and odour.” Sphagnum, along with Dicranum, are also used for lining diapers, in which it
keeps babies clean and warm. According to Gottesfeld and Vitt (1996), the famous “Johnson & Johnson
Company uses Sphagnum in diapers and sanitary napkins”. Most commonly utilised species for these
purposes is Sphagnum magellanicum. In Phillipines and New Guinea, women use mosses to decorate
ceremonial masks and also to decorate headwear and clothing (Tan, 2003). In some parts of Germany,
“wool was woven with Sphagnum to make good, cheap cloth”. Pant and Tiwari (1989) mentioned that
“women in villages of Kumaun, India, stuff mosses (Hylocomium, Hypnum, Trachypodopsis) into cloth
sacks to make head cushions”. Such cushions also absorb leaking water as they carry vessels.
BRYOPHYTES USED FOR CULTURING 32.12

Bryophytes, specially mosses, are particularly useful for special purposes, such as culturing or growing
of ferns (e.g. the moss Octoblepharum albidum) and orchids (e.g. species of Hypnum, Leucobryum,
Rhytidiopsis). Tan (2003) has mentioned that in Manila, “Leucobryum is substituted for peat moss and
induces good root sprouts on orchid cuttings”. Sphagnum has now proved to be an essential requirement
in the technique of air-layering. According to Pant (1989) Begonia and Fuchsia produce “buds and
flowers more profusely if their pot has a layer of moss to separate the humus-rich top and the bottom
soil”.
BRYOPHYTES AS SEED BEDS 32.13

Some bryophytes, specially mosses, are used as seed beds. According to Cox and Westing (1963),
“Sphagnum extracts induce germination of jackpine (Pinus banksiana) seeds, and aqueous extracts
of Polytrichum commune and Sphagnum spp. stimulate growth of Larix seedlings”. Mosses actually
provide the necessary humidity for germination. It has been proved by Equihua and Usher (1993) “that
Calluna vulgaris grew better and produced more flowers when it occurred in moss beds”.
MOSS GARDENS 32.14

In Japan, Great Britain, United States and some other developed countries, mosses are used to create
feelings of calmness, quietness or peace in gardens. These gardens are called moss gardens. Moss
gardens provide an uncluttered look of shades of green. In Japan, moss gardens are often associated
with Buddhist temples. One such temple, namely Kyoto’s Kokedera in Japan, is commonly called
the moss temple. Two most commonly used taxa for moss gardens belong to species of Pogonatum
and Polytrichum. Some other species grown in moss gardens are Dicranum scoparium, Leucobryum
bowringii, Rhizogonium dozyanum and Trachycystis microphylla. When fully grown, these mosses
provide looks of mounds or cushions, and look like miniature hills bearing rolling landscape. A well-
developed moss garden is at Chatsworth, Great Britain, where as many as 33 mosses and 4 liverwort
species have been grown to provide a calm, beautiful and peaceful atmosphere. Some beautiful mosses
grown in this garden are Dicranum scoparium, Hylocomium splendens, Neckera crispa, Polytrichum
commune and Rhizomnium punctatum. Ando (1972) has mentioned that a moss garden in the home of
famous poet William Wordsworth has cushions of Polytrichum commune. In spite of beautiful looks, a
moss garden is still an effort of wealthy people to increase the charm of their properties.
BRYOPHYTES AND GENETIC ENGINEERING 32.15

Much work has not been done so far on genetic engineering of bryophytes. Through the application
of the techniques of genetic engineering, however, bryophytes in the modern era are likely to be a
substantial source of human medicines, and provide a gene bank for making proteins, sugars, enzymes
or fatty acids. These techniques may also be soon helpful in permitting crop plants to survive cold,
drought, or infestations. With the genetic engineering techniques used for mosses, it is now at least
theoretically possible to manipulate the genomes of plants to endow them with the desirable traits
for human use. According to Hoffman (1992), some specific characteristics of bryophytes (e.g. their
ability to survive drought, and their ability to become functional within 24 hours) have aroused the
imagination of agriculturists from the point of view of genetic engineering.
Scott and Oliver (1994) have “isolated several genes specific for the recovery of desiccated
gametophytes of mosses”. Hohe et al. (2002) have used “the tiny moss (Physcomitrella patens) to
produce human proteins”. P. patens and some other mosses “have a high frequency of homologous
recombination”. It is the only bryophytic plant being used to produce the blood-clotting factor IX
for pharmaceutical purposes. Most mosses require no antibiotics during culture, and therefore, the
360 � Bryophyta
contamination of the final product can be avoided. The small size of Physcomitrella patens allows lab
culturing, thus reducing the possibility of escape of transgenetic plants.
1. Write a detailed essay on economic uses of bryophytes.

2. Bryophytes are now proving to be of definite medicinal uses for mankind. Comment in about
500 words.
3. Make a list of ten bryophytes of medicinal value for us.
4. Sphagnum is of great importance for us. Comment in about 300 words.
5. Write a note on an bio cs from bryophytes.
6. Bryophytes have some proven an -cancerous proper es. Comment.
7. Write short scien fic notes on:
(a) Bryophytes as source of food,
(b) Hor cultural uses of bryophytes.
8. Which moss is a proven bryophyte of the moss industry?
9. Bryophytes are now widely used as source of fuel and pes cides. Elaborate this statement.
10. What are moss gardens?
11. Two most commonly used bryophy c taxa for moss gardens belong to Pogonatum and
_____.
12. Write a note on bryophytes and gene c engineering.
Appendix I
Answers to TEST YOUR
UNDERSTANDING
Chapter 1
1. Braun (1864) 2. mosses 3. bryophytes 5. Bryophyta
6. No 10. Yes 12. flask
Chapter 2
2. Rothmaler (1951)
Chapter 3
1. Liverworts 3. Marchantiopsida 8. Only one (Takakia) 16. Haplomitrium
Chapter 4
1. Jungermanniales 3. only smooth-walled 4. Absent
5. Jungermanniales Anacrogynae 6. Jungermannineae 7. Metzgerineae
9. Jungermannineae 10. Madothecaceae 19. Jubula 27. endothecial
29. Cryptothallus
Chapter 5
1. Geothallus 2. Bottle hepatics 7. Aquatic 8. endothecial
9. very small
Chapter 6
2. Marchantiales 3. Schuster (1963) 4. Only one genus, namely Monoclea
6. Giant thallose-liverworts 8. Absent
Chapter 7
2. Chambered hepatics 3. Riccia 6. scales 7. tuberculate
8. not 9. dorsal 10. one 11. absent
13. Oxymitra 15. Riccia fluitans 16. terrestrial 23. Two
25. flask 29. Marchantia polymorpha 35. barrel
41. seta
362 � Bryophyta
Chapter 8
2. Proskauer (1957) 3. Horned liverworts 5. No 6. Absent
7. pyrenoid 8. intercalary meristem 9. amphithecial 10. Notothylaceae
11. Megaceros 16. Anthoceros 18. (c) 22. Embedded
Chapter 9
2. mosses 4. No 5. protonema 6. pleurocarpous
8. columella 9. absent 11. bog mosses 12. Andreaeales
15. No
Chapter 10
2. Sphagnales 3. Sphagnum 5. Only one, i.e. Sphagnum
6. perennial 17. amphithecial 22. Members of Andreaeales
25. Andreaea 26. Absent 27. Pseudopodium
29. Bryales 31. Funaria hygrometrica 32. No
39. microscopic 40. Four 41. Four-toothed moss 43. Archidium
Chapter 11
3. haploid 4. gametophyte 5. sporophyte, spores 6. Zoopsis argentea
7. foliose
Chapter 12
4. diploid 5. spores 6. foot 8. Riccia
10. capsule
Chapter 13
2. spermatophytes, sporophyte 3. meiosis 4. gametophytic
5. haploid 6. spore 8. homosporous 10. (a)
Chapter 14
5. day-neutral bryophytes 6. True 8. neutral
Chapter 15
4. heteromorphic 5. archegonia, bryophytes 7. W Hofmeister (1851)
10. Farlow (1874) 11. Pringsheim (1876) 13. Celakovsky (1874) 15. W H Lang (1909)
Chapter 16
2. phylogeny 3. genome 4. bryophytes 5. two
7. Anthocerotopsida 8. cladistics 10. (b) 11. Marchantiophyta
Chapter 17
6. Yes 7. phytochrome 8. metabolism 9. senescence
10. napthaline acetic acid, Trichlorophenoxy-acetic acid
Chapter 18
3. tuber 4. plural
Appendix I � 363
Chapter 19
1. archesporium 6. Pellia 7. Riccardia
8. amphithecium, endothecium 10. endothecial 11. amphithecial
Chapter 20
2. apogamy, apospory 4. Yes 10. Pringsheim (1876)
Chapter 21
3. ectohydric 4. Funaria hygrometrica 6. Ephemeropsis 8. Frullania
Chapter 22
4. Marchantia polymorpha 5. Marchantia polymorpha and Sphagnum
Chapter 23
3. Yes 7. bryometer
Chapter 24
3. Over 400 7. Marchantia polymorpha (Compound: marchantin-A)
8. Reboulia hemisphaerica
Chapter 25
9. Ribulose 1, 5-biphosphate carboxylase oxygenase
Chapter 26
3. S R Kashyap and P N Mehra
Chapter 27
2. Zygote 3. independent 4. gametophyte 5. spores
8. Riccia 9. Funaria 12. Reduction theory
Chapter 28
4. Leucobryum glaucum 5. external conduction
8. (d) 10. (d) 11. Polytrichales 13. food
Chapter 29
2. (b) 3. fresh 12. heteropycnotic 13. dioecious
17. allopolyploid, allopolyploidy
Chapter 30
2. autecology 6. habitat
Chapter 31
5. Psilophytales 8. All of these 15. fossil mosses
Chapter 32
8. Sphagnum 11. Polytrichum
Appendix II
Comparative Tables
Related to Bryophyta
Tables Already Discussed in the Text

1. Differences between algae and bryophytes (See Table 1.1; Chapter 1; Article 1.16.1, a)
2. Differences between bryophytes and pteridophytes (See Table 1.2; Chapter 1.;
Article 1.16.2, b)
3. Differences between Hepaticopsida, Anthoceropsida and Bryopsida (See Table 2.1;
Chapter 2; Article 2.2)
4. Comparison of sporogonium of Riccia and Marchantia (see Table 7.1; Chapter 7;
Article 7.8)
5. Differences between Anthocerotopsida and Hepaticopsida (See Table 8.1; Chapter 8;
Article 8.11)
6. Differences between Hepaticopsida and Bryopsida (See Table 12.1; Chapter 12;
Article 12.4.2)
7. Approximate number of spores per plant in some bryophytes (See Table 12.2; Chapter 12;
Article 12.5.4)
8. Origin, position and fate of archesporium in some common genera of bryophytes
(See Table 19.1; Chapter 19; Article 19.4)
9. Growth-forms classification for tropical forest bryophytes as proposed by Richards (1983)
(See Table 21.1; Chapter 21; Article 21.2)
10. Biological activity and effects of some liverworts and mosses of medicinal importance
(See Table 24.1; Chapter 24; Article 24.2).
Table A.1 Comparison of external features of gametophyte of Riccia, Marchan a, Pellia, Porella, Anthoceros, Sphagnum, Polytrichum and Funaria
S. NO. CHARACTER RICCIA MARCHANTIA PELLIA PORELLA ANTHOCEROS SPHAGNUM POLYTRICHUM FUNARIA
1. Plant form Thalloid, pros- Thalloid, prostrate, Thalloid, prostrate, Plant body is Thalloid, prostrate, Plant body has an Gametophores Plant body is foliose;
trate, dorsiven- flat, dorsiventral, flat and dorsiven- prostrate, flat, dorsiventrally flat erect and radially are differentiated gametophore is dif-
trally flattened, with a distinct tral; thalli with dorsiventral; and irregularly symmetrical gameto- into underground ferentiated into well-
with a distinct midrib; thalli larger sinuous overlapping contains a central lobed; midrib is phore which contains rhizome and many branched rhizoids,
midrib; each lobe than Riccia; an margins; larger than branched axis with not distinct. many green and radially sym- erect axis or stem
of thallus has an apical notch is Riccia but smaller two dorsal and one sessile leaves. metrical aerial, erect and many leaves
apical notch at the also present as in than Marchantia; ventral rows of branches containing
distal end, which Riccia. thinner than Riccia leaves. leaves.
contains grow- and Marchantia;
ing point of the with a distinct
thallus. midrib.
2. Branching Dichotomously Same as in Riccia. Same as in Riccia. Monopodial. Dichotomously Branches develop Branching is lateral; Same as in Polytri-
branched. branched, but from the axil of every branches are not chum.
thallus becomes fourth leaf; all axillary.
irregular due to branches are alike
unequal growth. in aquatic species,
but in semi-aquatic
species the branches
are of two types i.e.
pendent branches and
divergent branches.
3. Rhizoids ∑ Unicellular and ∑ Same as in Riccia ∑ Same as in Riccia ∑ Same as in ∑ Same as in ∑ Multicellular with ∑ Long, branched ∑ Long, well-
unbranched Riccia Riccia oblique septa multicellular, usu- branched, multicel-
∑ Present on ∑ Same as in Riccia ∑ Same as in Riccia ∑ Same as in ∑ Same as in ally coiled to form lular with oblique
ventral surface Riccia Riccia rope-like structure septa
in the midrib ∑ Present only on ∑ Present on under- ∑ Present only at
region. young gameto- ground rhizome the base of the
phores gametophore
∑ Two types, i.e. ∑ Same as in Riccia ∑ Only of one type, ∑ Same as in Pellia ∑ Same as in Pellia ∑ All are of only one ∑ All are of the same ∑ All are of the same
smoothwalled i.e. smoothwalled; type. type. type.
and tuberculate. tuberculate rhiz-
oids are absent
4. Scales Multicellular, Multicellular, Absent Absent Absent Absent Absent Absent
present in a single present in 2-3
row along the rows on either side
margins of the of midrib on the
thallus; all are of ventral surface; of
only one type. two types, i.e.
ligulate and
Appendix II �
appendiculate
365
Contd...
366
5. Leaves Absent Absent Absent Arranged in 3 Absent Thin, sessile, small- Leaves develop Leaves are sessile,
rows, i.e. 2 dorsal sized leaves present on underground simple, spirally
or lateral rows and in 3 vertical rows on rhizome and also arranged on the axis
one ventral row; the axis in 2/5 type on aerial parts; on and exhibit 3/8 type
leaves lack midrib; of phyllotaxy; midrib rhizome they show of phyllotaxy; each
they are unequally absent in mature 1/3 type of phyl- leaf possesses a clear
� Bryophyta
lobed. leaves. lotaxy but on aerial midrib.

branches leaves are
larger and show 3/8
type of phyllotaxy;
midrib present.
Table A.2 Comparison of the internal structure (anatomy) of the gametophyte of Riccia, Marchan a, Pellia and Anthoceros
S.NO. CHARACTER RICCIA MARCHANTIA PELLIA ANTHOCEROS
1. Differentiation Thallus is differentiated internally into Same as in Riccia Thallus is undifferentiated, i.e. Same as in Pellia
∑ upper photosynthetic region, and homogeneous
∑ lower storage region
2. Upper epidermis Not well-defined, but the terminal or Well-defined; wall of the Not well-defined Same as in Pellia
outermost cells of the upper photosynthetic upper epidermal cells appear
region function as epidermis. comparatively thick.
3. Air pores Not well-defined, but continuity of Well-developed, barrel-shaped air Absent Absent
epidermis is broken by some porelike pores, made up of 4-8 superimposed
structures, which are the outer openings tiers of cells; each tier has 4-5 cells;
of the air spaces present between air pores remain projected partly
photosynthetic filaments. above the thallus surface.
4. Photosynthetic Made up of vertical columns of Made up of many chambers; each Some dorsal cells of the middle All cells of the gametophyte contain
region chlorophyll-containing green cells; chamber contains some unbranched region, and also some cells of chloroplasts and are, therefore,
chloroplasts in these cells are discoid. or branched photosynthetic wings are green and represent photosynthetic.
filaments. photosynthetic region.
5. Pyrenoids Absent Absent Absent Each cell has a large chloroplast with a
single pyrenoid.
6. Air chamber Present in between the photosynthetic Many air chambers, arranged in Absent Absent
filaments. a single row, just below the upper
epidermis; a single-layered partition
wall separates them from each
other.
7. Storage region Ventral part of thallus is the storage region; Same as in Riccia; some mucilage All cells of thallus form storage Same as in Pellia; some mucilaginous
made up of parenchymatous cells. cells are also present. region. cavities are present in the ventral region.
8. Lower epidermis Possess smooth-walled and tuberculate Same as in Riccia; also possess Possess smooth-walled rhizoids. Same as in Pellia.
rhizoids. scales.
Table A.3 Comparison of male sex organs/antheridia of Riccia, Marchan a, Pellia, Porella, Anthoceros, Sphagnum, Polytrichum and Funaria
1. Differentia- Mostly monoecious; Mostly dioecious; Both monoecious as Mostly dioecious. Both monoecious Same as in Dioecious. Monoecious but
tion of sex in only a few dioecious. only a few monoe- well as dioecious. as well as dioe- Anthoceros. autoecious.
species cious. cious.
2. Position of Situated in the anthe- Present on the Present on the dorsal Develop on some Present in the an- Present singly in Present in groups Situated in clusters
antheridium ridial chamber, which lobes of peltate surface of the thallus specialised anthe- theridial chambers the axil of each at the base of each at the apices of an-
remains embedded in disc of the anthe- on either side of mid- ridial branches in covered by roof on leaf of specialised perichaetial leaf at theridial branches,
the dorsal surface of ridiophore. rib, embedded in the the axil of leaf. the dorsal surface male or antheridial the apices of male enclosed by a
thallus. antheridial chamber. of thallus. branches. gametophore. group of leaves.
3. Number of One antheridium Same as in Riccia. Same is in Riccia. Only one One to four One in the axil of Many antheridia Develop in groups
anthe- in each antheridial antheridum in or even more each leaf of the develop in groups at the terminal part
ridia in each chamber. the axil of each antheridia in each antheridial branch. at the tip of the of gametophore.
antheridial leaf of antheridial antheridial cavity. male gametophore.
chamber branche.
4. Structure Shortly-stalked, glo- Stalked, oval or Same as in Riccia; Stalk very long; Stalk multicellular, Stalk very long; Stalked, club- Massive stalk,
bose or club-shaped conical in shape; shape somewhat body spherical; body club-shaped; present in the axil shaped; jacket club-shaped body;
body, covered by a jacket single- spherical. antheridial jacket jacked single-lay- of leaf; jacket of cells surround single-layered
single-layered jacket. layered. 2-3 layered in the ered throughout. single-layered. mass of androgo- jacket; an opercu-
basal part. nial cells. lum is present at
the apex of jacket.
5. Antherozoids Unicellular, uni- Same as in Riccia. Spirally coiled, biflag- Rod-shaped, Same as in Riccia. Spirally coiled Spirally coiled, Same as in Polytri-
nucleate, curved and ellate. biflagellate. with a terminal uninucleate, and chum.
biflagellate. flagellophore, biflagellate.
bearing two
flagella.
Appendix II �
367
368
Table A.4 Comparison of female sex organs/archegonia of Riccia, Marchan a, Pellia, Porella, Anthoceros, Sphagnum, Polytrichum and Funaria
� Bryophyta
1. Position of Develop in archego- Develop on the Develop in groups of Develop at the Archegonia are Develop at the Develop at the Situated at the
archegonium nial chambers, em- dorsal surface of 4-10 on the dorsal sur- apex of archego- embedded on the apex of the arche- apex of arche- apices of female
bedded on the dorsal the disc present on face of thallus on either nial branch in large dorsal surface of gonial branches gonial head on branches inside the
surface of thallus. the top of arche- side of midrib. number between the thallus with in between a female plant; cluster of leaves;
goniophore. the leaves. only their cover perichaetial leaves, surrounded by intermingled with
cells projecting be- usually in groups coloured perichae- paraphyses.
yond the surface. of 2-5; position is tial leaves; appear
acrogynous. like a small flower.
2. Structure Flask-shaped struc- Same as of Riccia; Flask-shaped as in Same as of Pellia; Embedded in the Flask-shaped, Flask-shaped, Flask-shaped with
ture; shortly stalked; neck has 4-8 neck Riccia; jacket of neck neck is broad and thallus and are stalked structure stalked and greatly a multicellular
with a broad venter canal cells sur- consists of 5 vertical consists of 5 verti- in direct contact with a broad elongated; venter massive stalk, a
and long neck, venter rounded by 6 verti- rows of cells and cal rows of cells with the vegetative venter and long is many-celled broad venter and
wall is one-celled; cal rows of jacket encloses 6-8 neck canal and encloses 6-8 cells; neck canal twisted neck; thick; neck con- narrow twisted
neck consists of 6 cells; cover cells cells; venter is two- neck canal cells. cells are 4 in num- venter as well as sists of 6 vertical neck; wall of
vertical rows of cells, are indistinct. layer thick; cover cells ber; cover cells are lower portion of rows of cells; large venter is double-
6-9 cells in height and are 4 but indistinct. also 4. neck are 2-4 cells number of neck layered; 6 or more
possesses 4 neck ca- in thickness; neck canal cells are neck canal cells.
nal cells and 4 cover canal cells are 8-9 present; a single
cells; venter contains in number; cover cover cell is pres-
a ventral canal cell cells indistinct. ent at the top.
and an egg.
Table A.5 Comparison of the sporophyte of Riccia, Marchan a, Pellia, Porella, Anthoceros, Sphagnum, Polytrichum and Funaria
1. Parts of the Only capsule embed- Foot, seta and Foot, seta and capsule; Foot, seta and Foot, meristematic Foot and capsule; Foot, seta and Foot, seta and
sporophyte ded in the thallus; capsule; capsules horizontally placed on capsule; present zone and capsule; seta is very small capsule; present at capsule; develop at
and its posi- foot and seta absent. are seen in a disc the thallus. on the special- develop on the neck like sup- the apex of female the apex of arche-
tion of mature arche- ised archegonial dorsal surface of pressed tissue; gametophore; seta gonial branches.
goniophore, on the branches of female thallus; a distinct present at the is very long.
lower side. plants. seta is absent. apices of short,
bud-like female
branches.
2. Protective A double-layered Calyptra, Double-layered Calyptra, perianth Calyptra, also Calyptra and Calyptra surrounds Calyptra, which is
coverings calyptra around the perigynium and calyptra; and involucre known as invo- perichaetium; the young capsule shed at maturity.
young sporophyte. perichaetium. (formed by lucre. pseudopodium completely; at ma-
bracts of female develops in mature turity the calyptra
branches). sporophytes; covers only apical
calyptra and distal part of capsule.
end of pseudopodi-
um form a saclike
veginula.
3. Foot Absent Basal, bulbous part Conical, parenchyma- Indistinct foot; Bulbous, well- Well-developed, Dagger-shaped, Small, conical,
which anchors the tous, extends like a globose in young developed, paren- massive and parenchymatous. dagger-shaped
sporophyte to the collar around the base sporophyte chymatous, bulbous; haustorial structure.
disc of archegonio- of seta. in function.
phore.
4. Seta Absent. Present; short, Present; contains Same as in Pellia. Absent; the mer- Absent; a narrow, Very long, slender Long, slender,
stout, and connects abundant starch when istematic tissue, non-meristematic, reaching up to 5 twisted, and dif-
foot to the capsule; young. present in between necklike part, cm or more in ferentiated into
parenchymatous foot and capsule, however, is present length; differenti- epidermis, cortex
but its cells functions as seta. in between foot ated internally and central con-
become highly and capsule. into an epidermis, ducting strand.
vacuolated when sclerenchymatous
mature. hypodermis, par-
enchymatous cor-
tex and a central
cylinder composed
of hydroid cells.
5. Capsule Globose, embedded Oval. Same as in Marchantia. Oval Horn-like Globose. Polygonal. Obique or asym-
(i) Shape in the thallus. metrical; pear-
shaped; curved.
Appendix II �
Contd...
369
370
(ii) Wall Single-layered, which Single-layered Capsule wall com- Capsule wall con- Capsule wall is 4-6 Capsule jacket Capsule wall is Wall many-
also disorganises capsule wall; posed of 2 to 3 layers; sists of 2 to 6 cells layered, of which consists of 4-6 several-layered; layered; 2-3 inner
soon. persistent. persistent. in thickness. the outermost layer layers, of which all cells of wall wall layers are
forms epidermis, the outermost layer layers contain green and contain
ventilated with sto- is an epidermis chloroplasts. intercellular
mata; cells of inner containing rudi- spaces; 2-3 outer
� Bryophyta
wall layers contain mentary stomata. layers beneath

chloroplasts. the epidermis are
parenchymatous.
(iii) Arches- Originates from Same as in Riccia; Same as in Marchantia. Same as in March- Develops from Develops from Develops from Same as in Polytri-
porium endothecium; elaters elaters present. antia. the inner layer of inner layer of the outer layer of chum.
absent. amphithecium; amphithecium. endothecium.
pseudoelaters
present.
(iv) Elatero- Absent Absent Present Absent Absent Absent Absent Absent
phore
(v) Columella Absent Absent Absent Absent Present Present Present Present
(iv) Opercu- Absent Absent Absent Absent Absent Present Present Present
lum
(vii) Annulus Absent Absent Absent Absent Absent Present Present Present
(viii) Peris- Absent Absent Absent Absent Absent Absent Present Present
tome
6. Dehiscence of Spores liberate after Wall of the Wall of the capsule Same as in Pellia. Capsule dehisces Operculum sepa- Capsule dehisces Operculum sepa-
capsule death and decay of capsule splits splits longitudinally by longitudinal rates and spores and spores are rates and capsule
thallus without any longitudinally into into 4 valves; elaters splitting of its are dispersed by dispersed by dehisces due to
special mechanism. several valves; also help in spore walls into 1-4 special explosive Censor mechanism the hygroscopic
spore dispersal is dispersal. valves; dispersal is mechanism. with the help of movement of per-
facilitated also by facilitated also by epiphragm and istomial teeth.
the hygroscopic elaters, columella peristomial teeth.
nature of elaters. and valves of the
capsule.
Appendix III
Glossary of
Bryophytic Terms
Abaxial On the underside of a leaf, that is, pointing away from the stem
Acrocarpous Mosses, which have an upright stem, with the sex organs at the apex
Acrogynous Possessing the stem terminated by archegonia
Acropetal Development of organs successively towards the apex, i.e. the oldest at the
base and youngest nearest to the apex
Acuminate Possessing a gradually diminishing point
Adaxial On the top side of the leaf, that is, pointing towards the stem
Adnate United with another organ
Adventitious Produced abnormally
Alternation of The life cycle of bryophytes, pteridophytes, and spermatophytes, which
generations consists of a haploid gametophyte producing gametes followed by a diploid
sporophyte, producing spores
Alveolate Possessing cavities on the surface, generally of the spores
Amphithecium Outer layer of a developing sporophyte of a bryophyte
Anacrogynous Stem not being terminated by archegonia and continuing to grow
Androcyte Antherozoid mother cell; actually the cell which later develops into
antherozoid
Androgonial cell Any cell within an antheridium other than androcyte mother cell or androcyte
Annular Like a ring
Antheridium The male sex organs of the cryptogams
Antherozoid Unicellular, small, motile male gamete bearing flagella
(spermatozoid)
Antical Upper surface, usually of stem or leaf
Anticlinal A division perpendicular to the surface
372 � Bryophyta
Apogamy Asexual reproduction in which embryos and propagules are produced

without the occurrence of meiosis
Apophysis The sterile tissue at the basal part of the capsule of some mosses, present
immediately at the tip of seta and below the spore sac
Apospory Formation of a diploid gametophyte from the vegetative cells of the
sporophyte, that is, without the production of spores
Appressed Lying flat for the whole length of the organ
Archegoniatae Plants having archegonia
Archegonium The flask-shaped, female organ of bryophytes, pteridophytes and
gymnosperms; usually consists of a hollow neck, and a swollen base
containing the egg
Archesporium The first cell generation of sporogenous tissue, from which the spores of
sporogonium are ultimately derived
Areolate Small spaces marked on the surface, usually of spores
Autoecious Possessing male and female organs on the same plant but on separate
branches
Autotrophic Able to synthesize food from simple chemical compounds, using energy
from light or chemical reactions
Basipetal Such a development of organs in which the youngest structures are at the
base and oldest at the apex
Biseriate In two rows
Bisexual Organisms with male and female reproductive organs on the same
individual
Blepharoplast A cylindrical structure found at the base of cilia composed of nine sets of
triplet microtubules
Caducous Falling off early
Calyptra A protective covering around the young capsule derived from the
archegonium
Capillary Hairlike
Capsule Part of the sporogonium that contains spores
Carpocephalum Female receptacle
Chlorophyll Green colouring matter of plants
Chlorophyllous Containing chlorophyll
Chloroplasts Granules containing chlorophyll
Ciliate Fringed with hairs
Clavate Club-shaped
Columella Central column of sterile cells in a capsule
Appendix III � 373
Compressed Flattened out

Cryptogams A general name for all plants except gymnosperms and angiosperms;
cryptogams reproduce by spores
Costa Midrib of the thallus
Deciduous Falling off
Dehisce To split open
Dichotomous Dividing equally into two, especially of branches
Dimorphic An organism with two different forms
Dioecious With antheridia and archegonia on different plants
Diploid Of cells with two sets of chromosomes in their nuclei; the sets are set to be
homologous
Diploid generation The sporophyte
Distichous Disposed in two rows
Divergent Spreading apart
Dorsal The surface of the leaf, away from the stem; the upper surface of a thallus
Dorsiventral With dorsal and ventral surfaces
Egg A female gamete
Elaters Bunch of long, thin cells in the capsule of the sporophyte of a liverwort;
have spiral thickening of the cell wall; alter their position with changes in
humidity, and help in dispersal of spores
Embryo The young plant developed within the archegonium; the product of the
repeated mitotic divisions of the zygote
Embryophyta A group that includes bryophytes, pteridophytes and spermatophytes
Endemic Of taxa that are found only in one particular place or area
Endogenous Arising from deep-seated tissue
Endothecium The inner tissues in the young sporophyte of bryophytes, giving rise to the
sporogenous tissue and/or the columella
Epibasal Forming the upper part of the embryo
Epiphragm A membrane which closes the opening of theca in the sporophyte of some
mosses, e.g. Polytrichum
Epiphyllous Developing on leaves
Exogenous Arising from the superficial tissue
Exotic Non-native
Extant Species which exist at present
Extinct Species which no longer exist
Fertilization Fusion of a male gamete with a female gamete to form a zygote
374 � Bryophyta
Filiform Threadlike
Fimbriate Fringed
Flagellum A fine thread-like branchlet; a long motile thread, consisting of a membrane
enclosing a series of parallel microtubules
Foliar Belonging to a leaf
Foliose With leaves
Foot An organ of attachment and nutrition; the lowest part of the sporophyte
Fossil The remains or marks left by dead organisms, converted to stone over
geological time
Frond The leaf of a fern or palm
Fusiform Tapering at both ends like a spindle
Gametangium Any organ which produces gametes
Gamete A haploid sex cell whose function is to join with a gamete of the opposite
sex, to form a diploid zygote
Gametophyte The haploid generation in an alternation of generations; in bryophytes, the
gametophyte is the main vegetative stage; the gametophyte is the generation
producing gametes
Gemmae Small groups of green cells produced in the cup-shaped structures on the
surface of thalloid liverworts
Gemmiferous Bearing gemmae
Generation A set of individuals of roughly equal age or stage of development; the parents
are one generation, and the progeny represents the next generation
Granulose Composed of grains
Habit The appearance of a plant, e.g. herb, shrub or tree
Habitat The place or kind of place in which an organism, community or association
is found
Haploid The cells with one set of chromosomes in their nuclei
Haploid generation The gametophytic generation
Hepatic A liverwort
Heteromorphic A condition in which two phases of the life-cycle are morphologically
dissimilar
Heterosporous Producing two kinds of spores, usually of different sizes
Homologous Of two similar chromosomes which pair with each other during meiosis
Hyaline Transparent, or without colour
Hygrophyllous Plants which require a large supply of moisture for their growth and development
Hypobasal Forming the lower part of the embryo
Hypogynous Inserted below the archegonium

IAA Indole acetic acid, the most common auxin
Incised Cut sharply
Incubous Of a kind of growth in leafy liverworts, in which the front of each leaf lies on
top of the leaf in front of it
Innovation A newly-formed shoot which continues growth at the death of the older
stem
Intine The inner coat of the wall of a spore
Involucre A tubular structure serving to protect the archegonia and calyptra
Involute Having the edges rolled inwards
Juvenile Very early forms
Karyogamy Fusion of two nuclei after plasmogamy
Keel A ridge-like the keel of a boat
Lacerate Irregularly torn or cleft
Lacuna A space, particularly in between the tissues
Lamella A plate of tissue
Lamina The parts of the leaf on either side of the midrib
Leafy liverwort A liverwort in which the gametophyte has a simple stem, growing from the
apex and bearing small leaves in rows along it
Lenticular Like a double convex lens
Lignified Woody
Liverwort A group of bryophytes, which differ from mosses in having less differentiated
cells in the gametophyte, and in usually having elaters in the capsule
Lumen The cavity inside a cell
Lysigenous A cavity developed by the disintegration and dissolution of cells
Marsupium The fruiting receptacle of certain liverworts
Medulla The parenchyma or sclerenchyma inside the vascular strands
Meiosis The cell division that produces haploid sex cells from diploid cells; also
called reduction division
Meristem Any tissue of actively dividing cells, which produce the cells of the other
plant tissues
Micron A measure of length equal to one millionth of a metre (or 1/1000 millimetre);
symbol µ
Midrib The costa (thallus); vein of the leaf
Mitosis Vegetative or somatic cell division in which chromosomes in the nucleus are
duplicated into two chromatids
376 � Bryophyta
Monoecious A condition when antheridia and archegonia develop on the same plant
Morphogenesis The development of shape and structure of organs and tissues
Moss One of the three main groups of bryophytes; mosses differ from liverworts
in having more differentiated cells in the gametophyte; the gametophyte of a
moss usually possesses stem with leaves
Multifid Divided into many lobes
Neck The upper part of the archegonium
Neck canal cells The cells present in the neck canal of an archegonium
Nodulose Knotted, or thickened
Obcordate Inversely heart-shaped
Ontogeny The process of development of an individual from zygote to adult
Oogamous Possessing a small motile male gamete and a large nonmotile female gamete,
as in bryophytes and pteridophytes
Oogonium A reproductive organ, which produces female gametes
Oosphere The female gamete, produced in an oogonium
Oospore A dormant, thick-walled zygote, formed after the fertilization of an oosphere
Operculum A lid, which covers the pore at the apex of a moss capsule; the operculum
opens to allow spores to escape
Ostiole The tubular neck of the cavity containing antheridia
Palea The chaff scales or ramenta of many ferns
Papillose Covered with papillae
Paraphyllia Very fine, minute leaflike appendages on the stem of mosses and some
liverworts
Paraphyses Small, one-cell-thick hairs, often with a large round cell at the top; grow
among the antheridia in mosses
Paroicous A condition in mosses when antheridia are present below the archegonia on
the same branch or stem
Parthenogenesis A condition in which an embryo develops from an unfertilized egg
Peat A kind of litter layer found in some very wet or waterlogged habitats, such
as bogs, which decomposes very slowly, often under very acidic conditions;
peat layers may be as thick as several layers
Pedicel A short stalk
Perianth Inflated envelope surrounding the fertilized archegonium
Perichaetial Leaves surrounding the archegonia
Perennation The survival of an organism over successive years, or of a dormant organ
during unfavourable seasons
Periclinal A cell division, in which wall runs parallel to the surface of a plant organ
Perigonium Some specialised bracts around the male flower or antheridia
Perigynium An involucre of female inflorescence in some bryophytes
Peristome A set of tooth-like plates under the operculum of the capsule of a moss; the
plates are called peristomial teeth
Phloem One of the conducting tissues in the vascular system; phloem, unlike xylem,
is mainly a living tissue made up of sieve elements and companion cells
Phylogeny The evolutionary history of an organism or taxonomic group of organisms
Pleurocarpous Of mosses, which have a stem with many branches, spreading across the
ground; sex organs are borne on short side branches
Postical Belonging to lower surface of thallus, stem or leaf
Procumbent Lying along the ground
Propagule Any reproductive unit which gives rise to a new individual
Protandrous When male sex organs are produced prior to the formation of female sex
organs
Protonema The young, often filamentous, gametophyte of a moss in the early stages
after the germination of the spore
Pseudoelaters Sterile cells mixed with spores in the capsule of Anthoceros
Pseudoperianth A soft gametophytic sheath, which usually develops late and surrounds the
young sporogonium in some bryophytes
Pyrenoid A small grain of protein in the chloroplast of some plants, around which
starch is deposited
Radial Spreading from a common axis or centre
Receptacle The top of the stalk bearing reproductive organs (antheridia or archegonia)
Regeneration The growth of new tissue on a part of a plant that has been damaged; or, the
growth of new plants from perennating organs, e.g. rhizome
Reniform Kidney-shaped
Reproduction The process in which an organism produces offspring like itself
Reticulate Like a network
Retort cells Specialised enlarged cortical cells of the young branches of some bryophytes,
e.g. Sphagnum
Rhizoid A thread-like cell which grows from the lower surface or base of a bryophyte,
and functioning like a root
Rhizome A stem which grows along under the ground, bearing buds which produce
shoots; a root-like underground stem
Rosette Structure in which leaves or young parts are arranged in a light spiral; or a
pattern of organs like a rose
378 � Bryophyta
Saccate Like a bag

Scale A flat, thin, semitransparent plate of cells
Schizogenous Internal spaces which are produced by separation of cells
Sclereid Kind of cell found in the sclerenchyma of some plants, with heavily lignified
walls
Serrate Toothed like a saw
Sessile Without a stalk
Seta Stalk of the sporophyte of a bryophyte
Sinuate Wavy
Somatic Of any process or part of an organism that is not connected with sexual
reproduction, e.g. mitosis is a somatic cell division
Sperm Motile, flagellated, male sex cell
Spore Small round cell with a thick wall from which a whole new plant is produced
in bryophytes, pteridophytes and spermatophytes; spores are haploid and are
produced by the sporophyte
Spore mother cell A cell which divides by meiosis to produce spores
Sporogenous A tissue in which spores are produced
Sporogonium The spore-bearing generation developing from the fertilized egg; the
sporophyte of bryophytes
Sporophyte The non-sexual generation, or the part bearing spores; the diploid generation
in an alternation of generations
Stellate Star-shaped
Stylus A small, awl-like lobule
Synoicous A condition in mosses, when the archegonia and antheridia are mixed
together in the same involucre
Taxon Any taxonomic group, e.g. a species, a genus, a family
Terete Cylindrical, not angular
Tetrad (Tetrahedral) Spores remaining united in groups of four until mature
Thalloid (Thallose) Possessing the form of a thallus
Thallus More or less undifferentiated plant body, flat and broad like a frond, and
without distinct roots, stems and leaves
Theca Major part of capsule of mosses
Trigonous Having three obtuse angles
Trilete Bearing a triradiate tetrad scar
Tristichous Arranged in three rows
Truncate Cut abruptly
Tuber An organ of perennation and vegetative reproduction, a tuber is a thick

underground stem in which food remains stored; buds on the tuber grow into
new plants
Turgor The tension on a cell wall due to the pressure of water inside the cell
Vacuole A liquid-filled space in a cell, surrounded by a membrane
Vascular cylinder A tube of vascular tissue consisting of xylem and phloem
Variety A taxonomic group within a species or a subspecies
Vegetative Any part of a plant which is not involved in sexual reproduction
Vegetative A type of asexual reproduction in which a whole new plant is produced from
reproduction an organ, which is not involved in sexual reproduction
Vein One of the many lines which can be seen on the surface of a leaf, marking the
position of a vascular bundle
Venter The lower part of the archegonium
Ventral Pertaining to that surface of a flattened thalloid plant which faces the
substrate
Ventral canal cell A cell in the upper part of the venter of an archegonium and situated above
the oosphere
Vermiform Worm-shaped
Verrucose Covered with wart-like protuberances
Vesicle Any small body in a cell or organelle which is surrounded by a membrane
and contains products of metabolism
Vestigial A small or reduced structure
Wavelength The length of a wave of light; different wavelengths have different colours
and different levels of energy
Weed A plant growing where it is not wanted by humans
Whiplash flagellum A flagellum without hairs on its surface
Whorl A group of three or more plant structures arising at the same level on a stem
and forming a ring around it
Xeromorphic Plants with characteristics suited to very dry habitats, such as deserts
Xerophyte A plant that lives in a desert or other dry habitats
Xylem A tissue in the vascular system of a plant, consisting of tracheids, vessels,
parenchyma and sclerenchyma
Zoospore A motile flagellated spore
Zygote A diploid cell which is produced by the fusion of two haploid gametes
Zygotene A stage in the first meiotic prophase, when the homologous chromosomes
come together to form bivalents
Appendix IV
Globally Published
Books and Some Selected
Theses on Bryophyta
Arnell, S; 1956; Illustrated Moss Flora of Fennoscandia, 1, Lund.

Asakawa, Y; 1983; Chemical Constituents of the Hepaticae, Academic Press, London.
_____ 1990; Bryophytes: Their Chemistry and Chemical Taxonomy, Oxford University Press, Oxford.
Barkman, J J; 1958; Phytosociology and Ecology of Cryptogamic Epiphytes, Assen.
Bates, J W and Farmer, A M; 1992; Bryophytes and Lichens in a Changing Environment, Clarendon Press,
Oxford.
Bopp, M; 1983; ‘Developmental Physiology of Bryophytes’; In: New Manual of Bryology, Hattori Botanical
Laboratory, Japan.
Bower, F O; 1908; The Origin of Land Flora, London.
Brothers, V F; 1925; ’ Musci’; In: Engler and Prantl Die Naturlichen Pflanzenfamilien; Volumes 10 and
11, Leipzig.
Brown, D H; 1992; ‘Impact of Agriculture on Bryophytes’; In: Bryophytes and Lichens in a Changing
Environment (ed. J W Bates and A M Farmer), Clarendon Press, Oxford.
Burgeff, H; 1943; Genetische Studien an Marchantia, Gustav-Fisher, Jena.
Campbell, D H; 1918; The Structure and Development of Mosses and Ferns, Macmillan, London.
_____ 1940; The Evolution of Land Plants (Embryophyta), Stanford University Press, California.
Cavers, F; 1911; The Inter-relationships of the Bryophyta, New Phytology Report No. 4, Cambridge.
Chopra, R N; 1981; ‘Some Aspects of Morphogenesis in Bryophytes’; In: Recent Advances in Cryptogamic
Botany, Palaeobotanical Society, Lucknow.
_____ and Kumra, P K; 1988; Biology of Bryophytes, Wiley Eastern Limited, New Delhi.
_____ and Bhatia, S C; 1990; Bryophyte Development: Physiology and Biochemistry, CRC Press, Boca
Raton.
Clarke, G C S and Duckett, J G; 1979; Bryophyte Systematics, Academic Press, New York.
Cove, D J and Ashton, N W; 1984; The Experimental Biology of Bryophytes, Academic Press, New York.
Crandall, S B and Stotler, R E; 2000; ‘Morphology and Classification of the Marchantiophyta’; In: Bryophyte
Biology (ed. A J Shaw, et al.), Cambridge University Press, Cambridge.
Appendix IV � 381
Crum, H; 2001; Structural Diversity of Bryophytes. University of Michigan Herbarium, Ann Arbor.
_____ and Anderson, L E; 1981; Mosses of Eastern North America, California University Press, New
York.
Dixon, H N; 1924; The Student’s Handbook of British Mosses, London.
_____ 1932; ‘Classification of Mosses’; In: Verdoorn’s Manual of Bryology, The Hague.
Doyle, W T; 1970; The Biology of Higher Cryptogams, Macmillan, London.
Dua, S; 1983; Morphogenetic and Physiological Studies on Some Bryophytes, PhD Thesis, University Of
Delhi, India.
Duckett, J G and Renzaglia, K; 1991; In: Heidelberg Sympsoium on Physiology, Developmental Cell Biology
and Genetics of Bryophytes.
Dyer, A F and Duckett, J G; 1984; The Experimental Biology of Bryophytes, Academic Press, New York.
Eschrich, W; 1975; ‘Bidirectional Transport’; In: Phloem Transport (ed. S Aronoff et al. ), Plenum Press,
New York.
Flowers, S; 1973; Mosses: Utah and the West, Brigham Young University, Proto, Utah.
Fogg, G E; 1998; The Biology of Polar Habitats, Oxford University Press, New York.
Fritsch, R; 1982; Index to Plant Chromosome Numbers: Bryophyta, Junk, The Hague.
Gangulee, H C; 1969–1980; Mosses of Eastern India and Adjacent Regions, Kolkata.
Geiger, H; 1990; In: Bryophytes: Their Chemistry and Chemical Taxonomy, Clarendon Press, Oxford.
George, S; 1997; Moss Gardening, Timber Press, Portland, OR.
Gersen, U; 1982; ‘Bryophytes and Invertebrates’; In: Bryophyte Ecology (ed. J E Smith), Chapman and Hall,
New York.
Glime, J M and Saxena, D K; 1990; Uses of Bryophytes, Today and Tomorrow Publishers, New Delhi.
Goebel, K; 1905; Organography of Plants Part II, Hafner Publishing Company, London.
Goffinet, B, Hollowell, V and Magill, R; 2004; Molecular Systematics of Bryophytes, Missouri Botanical
Garden Press, St. Louis.
Graham, L E; 1993; The Origin of Land Plants, Wiley, New York.
Hartmann, E and Weber, M; 1990; In: Bryophyte Development: Physiology and Biochemistry, CRC Press.
Hebant, C; 1977; The Conducting Tissues of Bryophytes, J Cramer, Germany.
_____ 1979; ‘Conducting Tissues in Bryophyte Systematics’; In: Bryophyte Systematics (ed. G C S Clarke
and J G Duckett), Academic Press, London.
Hedderson, T A and Cox, C J; In: 1998; In: Bryology For the Twenty-first Century (ed. J W Bates), British
Bryological Society and Maney Publishing, Leeds.
Hughes, J G; 1955; Physiology of Sexual Reproduction in Bryophytes, PhD Thesis, University of Wales,
Cardiff.
Huneck, S; 1983; ‘Chemistry and Biochemistry of Bryophytes’; In: New Manual of Bryology (ed. R M
Schuster), The Hattori Botanical Lab, Japan.
Ignatov, M S and Ignatova, E A; 2003; Moss Flora of the Middle European Russia, KMK Scientific Press
Ltd, Moscow.
Kapur, A; 1983; In Vitro Studies on Some Bryophytes, PhD Thesis, University of Delhi, India.
Karunen, P; 1990; In: Bryophytes: Their Chemistry and Chemical Taxonomy, Clarendon Press, Oxford.
382 � Bryophyta
Kashyap, S R; 1929; Liverworts of the Western Himalayas and the Punjab Plains, Part I and II, Punjab
University Publication, Lahore.
Kimmerer, R W; 2003; Gathering Moss: A Natural and Cultural History of Mosses, Oregon State University
Press, Corvallis, OR.
Knoop, B; 1984; In: The Experimental Biology of Bryophytes, Academic Press, London.
Kumar, S S; 1983; ‘Cytogenetics of Indian Bryophytes’; In: Genetical Research in India, ICAR, New
Delhi.
Kumra, P K; 1977; Experimental Studies on Some Delhi Mosses, MSc Thesis, University of Delhi, India.
_____ 1981; Morphogenetic and Physiological Studies on Some Mosses, PhD Thesis, University of Delhi,
India.
Lawton, E; 1971; Moss Flora of Pacific Northwest, Hattori Botanical Lab, Japan.
LeBlanc, F and Rao, D N; 1975; ‘Effects of Air Pollution on Lichens and Bryophytes’; In: Responses of
Plants to Air Pollution (ed. J B Mudd), Academic Press, New York.
Longton, R E; 1988; The Biology of Polar Bryophytes and Lichens, Cambridge University Press,
Cambridge.
_____ and Schuster, R M; 1983; In: New Manual of Bryology, Hattori Botanical Lab, Japan.
Luttge, U and Higinbotham, N; 1979; Transport in Plants, Springer-Verlag, New York.
Macvicar, S M; 1926; The Student’s Handbook of British Hepatics, London.
Malcolm, B and Malcolm, N; 2000; Mosses and Other Bryophytes: An Illustrated Glossary, Micro-optics
Press, New Zealand.
Markham, K R and Porter, J G; 1978; ‘Chemical Constituents of the Bryophytes’; In: Progress in
Phytochemistry (ed. L Reinhold et. al.), Pergamon Press, New York.
Matzke, E B and Randzens, L; 1969; ‘Apospory in the Hepaticae’; In: Current Topics in Plant Sciences (ed.
J E Gunckel), Academic Press, New York.
Mishler, B D; 1988; ‘Reproductive Ecology of Bryophytes’; In: Plant Reproductive Ecology (ed. A J Lovett),
Oxford University Press, London.
Newton, A E and Tagne, R S; 2007; Pleurocarpous Mosses: Systematics and Evolution. CRC Press, Boca
Raton.
Newton, M E; 1979; In: Bryophyte Systematics, 207, Academic Press, London.
_____ 1984; In: Experimental Biology of Bryophytes, 65, Academic Press, London.
_____ 1989; Practical Guide to Bryophyte Chromosomes, British Bryological Society, Cardiff.
Niklas, K J; 1997; The Evolutionary Biology of Plants, University of Chicago Press, USA.
Nilsson, E and Bendz, G; 1973; ‘Flavonoids in Bryophytes’; In: Chemistry in Botanical Classification (ed.
G Bendz), Academic Press, New York.
Paolillo, D J Jr; 1974; ‘Motile Male Gametes of Plants’; In: Dynamic Aspects of Plant Ultrastructure (ed. A
W Robard), McGraw-Hill, London.
Parihar, N S; 1987; An Introduction to Embryophyta Vol. 1, Bryophyta, Central Book Depot, Allahabad.
Proctor, M C F; 1979; ‘Structure and Ecophysiological Adaptations in Bryophytes’; In: Bryophyte Systematics
(ed. G C S Clarke et al. ), Academic Press, London.
_____ 1981; ‘Physiological Ecology of Bryophytes’; In: Advances in Bryology (ed. W J Schultz-Motel), J
Cramer, Germany.
Appendix IV � 383
Puri, P; 1973; Bryophytes: A Broad Perspective, Atma Ram and Sons, New Delhi.
_____ 1986; Bryophytes: Morphology, Growth and Differentiation, Atma Ram and Sons, New Delhi.
Rao, D N; 1982; ‘Responses of Bryophytes to Air Pollution’; In: Bryophyte Ecology (ed. A J E Smith),
Chapman and Hall, London.
Rashid, A; 1968; Morphogenetic Studies on Some Indian Bryophytes, PhD Thesis, University of Delhi,
India.
_____ 1998; An Introduction to Bryophyta: Diversity, Development and Differentiation, Vikas Publishing
House Pvt. Ltd., New Delhi.
Rawat, M S; 1976; Morphogenetic Studies on Some Bryaceae, PhD Thesis, University of Delhi, India.
Reimers, H; 1954; ‘Bryophyta’; In: Engler’s Syllabus Der Pflanzenfamilien.
Renzaglia, K S and Vaughan, K C; 2000; ‘Anatomy, Development and Classification of Hornworts’; In: The
Biology of Bryophytes (ed. J Shaw and B Goffinet), Cambridge University Press, Cambridge.
Richards, P W; 1959; ‘Bryophyta’; In: Vistas of Botany (ed. W B Turrill).
_____ 1983; ‘The Ecology of Tropical Forest Bryophytes’; In: New Manual of Bryology, Vol. 2(ed. R M
Schuster), Hattori Botanical Lab, Japan.
Richardson, D H S; 1981; The Biology of Bryophytes, Blackwell Science Publication, Oxford, U K.
Sainsbury, G O K; 1955; A Handbook of New Zealand Mosses, Wellington, New Zealand.
Scagel, R F; 1982; Non-vascular Plants: An Evolutionary Survey; Wadsworth.
Schofield, W B; 1985; Introduction to Bryology, Macmillan, New York.
_____ 2002; Field Guide to Liverwort Genera of Pacific North America, Global Forest Society, University
Of Washington Press, Seattle, USA.
Schuster, R M; 1966; The Hepaticae and Anthocerotae of North America, Columbia University Press,
New York.
_____ 1984; New Manual of Bryology, Vol. 1 and 2, The Hattori Botanical Lab, Japan.
Schlesinger, W H; 1997; Biogeochemistry: An Analysis of Global Changes, Academic Press.
Schwabe, W W; 1990; In: Bryophyte Development—Physiology and Biochemistry, CRC Press.
Scott, D H; 1955; Introduction to Structural Botany -I, Flowerless Plants, London.
Seabury, F; 1975; Sporogenesis in Selected Genera of Musci, PhD Thesis, Texas University, Texas.
Shaw, J A and Goffinet, B; 2000; Bryophyte Biology, Cambridge University Press, New York.
Shlyakov, R N; 1975; The Liverworts: Morphology, Phylogeny and Classification, Navka, Leningrad (in
Russian).
Siegel, S M; 1962; The Plant Cell Wall, Pergamon Press.
Singh, D K; 2002; Notothylaceae of India and Nepal, Bishen Singh, M P Singh, Dehradun, India.
Smith, A J E; 1978; Moss Flora of Britain and Ireland, Cambridge University Press, Cambridge, UK.
_____ 1982; Bryophyte Ecology, Chapman and Hall, London.
Smith, G M; 1955; Cryptogamic Botany, Vol. II, Bryophytes and Pteridophytes, McGraw-Hill, New York.
Smith, H; 1975; Phytochrome and Phytomorphogenesis, McGraw-Hill, London.
Sood, S; 1972; Experimental Studies on Some Indian Mosses, PhD Thesis, University of Delhi, India.
Tiwari, S D; 1984; Studies on Bryophytes of Kumaon Himalayas, PhD Thesis, Kumaon University, Nainital,
India.
384 � Bryophyta
Udar, R; 1970; An Introduction to Bryophyta, S M Prakashan, Lucknow.

_____ 1976; Bryology in India, Chronica Botanica Co., New Delhi.
Vashistha, B D; 1985; In Vitro Investigation on Some Indian Bryophytes, PhD Thesis, University of Delhi,
India.
Vashistha, B R; 1991; Botany for Degree Students: Bryophyta, S Chand and Co. Ltd, New Delhi.
Verdoorn, Fr; 1932; Manual of Bryology, The Hague.
Vitt, D H; 1984; ‘Classification of Bryopsida’; In: New Manual of Bryology (ed. R M Schuster), Hattori
Botanical Lab, Japan.
Watson, E V; 1955; British Mosses and Liverworts, Cambridge.
_____ 1971; The Structure and Life History of Bryophytes (3rd ed.), Hutchinson University Library,
London.
Watson, H; 1947; Woodland Mosses, Forestry Commission Booklet No. 1, London.
Willson, M F and Burley, N; 1983; Mate Choice in Plants, Princeton University Press, Princeton, New
Jersey.
Wyatt, R and Anderson, L E; 1984; In: Experimental Biology of Bryophytes, Academic Press, London.
Zyrd, J P and Schaefer, D; 1999; In: Heidelberg Symposium: Physiology Developmental Biology, Cell
Biology and Gametes of Bryophytes, 29.
Index
A Alternation of generations 10, 170

Amino acids 285
Abscisic acid 282 Amphidium 338
Accessory chromosomes 331, 336 Amphigastria 35, 116
Acetogenins 293 Amphithecial 19, 94
Acid rain 288 Amphithecial in origin 158
Acrocarpous 140, 99 Amphithecium 43, 116, 136, 137
Acrocarpous mosses 16, 142, 247 Amylom layer 192
Acrocladium 342 Anabaena 287
Acrogynae 35 Anacrogynae 35
Acrogynous 29, 96 Anacrogynous 31, 104
Acrogynous jungermanniales 37, 161 Andreaea 18, 100
Acromastigum 202 Andreaeaceae 163
Acroschisma 163 Andreaeales 141
Actinothuidium 357 Andreaeobryopsida 20, 129
Active dehiscence 213 Andreaeobryum 163
Adventitious branch 7, 88, 258 Andreaeopsida 20
Affinities 12 Androcyte mother cells 89
Affinities of anthocerotales 135 Androcytes 102, 146
Affinities of bryophytes 12 Androgonial cells 102
Affinities of Sphagnum 161 Aneura 60
Age of bryophytes 4 Annulus 148, 343
Air chambers 83, 96, 141, 232, 233 Anoectangium 344
Air-gun mechanism 159, 312 Anomodon 281, 356, 357
Air pore 86, 98, 311, 103 Antheridial branch 154
Air space 178 Antheridial chambers 101
Aitchinsoniella 81 Antheridial initial 170
Algae 13, 14, 90, 103 Antheridiophore 96, 101
Algal ancestors of bryophytes 348 Antheridium 7, 155
Alkanes 282 Antherozoid 55, 90, 177
Allergy-inducing compounds 296 Anthoceropsida 1, 253, 169
Allopolyploid 337 Anthoceros 7
Allopolyploidy 337 Anthocerotaceae 117
Alternaria 297 Anthocerotae 2, 269
386 � Index
Anthocerotalean origin 135 Assimilatory filaments 311

Anthocerotalean origin of pteridophytes 135 Asterella 7
Anthocerotales 117, 234 Athalamia 81
Anthocerotales: A synthetic group 137 Atracheata 5
Anthocerotes 16 Atrichum 211, 354, 355
Anthocerotophyta 3 Atrium 335
Anthocerotopsida 17, 273, 156 Aulacomnium 254, 355
Anthocyanins 283 Autecology 341
Anthorhyniaceae 138 Autoecous Mosses 144
Antibacterial activities 281 Autopolyploid 337
Antibacterial activity 297 Autopolyploidy 337
Antibiotic activity of bryophytes 281 Auxin 230, 282
Antibiotics from bryophytes 355 Axis cladia 7
Antical lobe 37
Anti-cancerous activity 355 B
Antifungal property 281
Bajdaievia 351
Antimicrobial activity 281
Barbella 342, 355
Antiobesity activity 300
Barbula 143, 210, 229, 20, 354, 355
Antithetic 233
Bartramia 208, 228, 356
Antithetic theory 234
Bartramidula 227, 19, 123
Antitumor elements 281
Bazzania 199, 289, 245
Antiviral activity 355
Bicyclohumulenone 295
Aongstroemia 5
Biogeochemical 302
Apical cell 87
Bioindicators 286, 102
Apical growth 120
Biological Fixation of Nitrogen 307
Apogamous sporophytes 271, 272
Biologically 294
Apogamy 11, 234
Biologically active compounds 293
Apogamy and apospory 269
Biomonitors 289
Apolar 109
Blasia 201, 216, 287
Apophyses 176
Blasiaceae 52
Apophysis 145, 175, 321
Blepharoplast 89
Apospory 7, 11, 312
Blepharostoma 203
Apospory in liverworts and hornworts 274
Bog 141
Appendiculate scale 96, 97
Bog moss 150, 151
Archegonial branches 155, 172
Books 380
Archegonial head 192
Bottle-hepatics 70
Archegonial initial 156
Brachythecium 207
Archegoniatae 233
Bracket mosses 277
Archegoniophore 96, 103
Bracts 38
Archegonium 7, 125
Bryales 141
Archegonium development 42
Bryidae 166
Archeogoniophore 82
Bryobionta 81
Archesporium 23, 74
Bryobiotina 3, 24
Archidiaceae 185
Bryokinin 282
Archidiales 185
Bryometer 288
Archidium 185
Bryophyta 1, 3, 4
Archilejeunea 221
Bryophyte phylogeny 243
Arnellia 212
Bryophytes 1, 3, 13
Aromatic compounds 293
Bryophytes and genosystematics 244
Aspergillus 297
Bryophytes as food source 355
Aspiromitus 117
Index � 387
Bryophytes as indicators 286 Central strand 199

Bryophytic community 341 Cephalozia 212, 236
Bryophytic terms 371 Cephaloziella 81
Bryopsida 1 Ceratodon 222, 341
Bryopteris 7 Ceratolejeunea 221
Bryum 181 Cetraria 308
Buchtia 351 Chalcones 283
Bug mosses 169 Chambered hepatics 147
Bulbil 5, 259 Chemotropism 280
Buxbaumia 181 Chiloscyphus 297
Chloronema 147
C Chloronemal branches 13
Caenorphabdiitis 297 Chlorophyceae 120
Callicostella 203, 12 Chloroplast 203
Calliergon 287 Chondrophyllum 2
Callus 270 Chondrus 2
Calobrium 30 Chorisodontium 305
Calobryaceae 29 Chromatin 332
Calobryales 28, 57 Chromosome numbers 333
Calobryum 24, 292, 63, 127 Cinclidotus 291
Calymperes 289, 133 Cladonia 17
Calypogea 337 Cladothamnus 344
Calypogeia 5, 281 Classification 16, 19
Calyptra 9, 273 Classification of bryophytes 23
Camptothecium 342 Classification of hepaticopsida 200
Campylopus 258, 297 Cleaning agents of toxic waste 292
Camtothecium 278 Cleistocarpous 214
Candida 281, 183 Climacium 356
Canopy-formers 342 Climacium dendroides 2
Caphaloziella 207 Club-moss 63
Capsule 6, 57, 180, 183 Coleochaete 234, 312
Capsule of Funaria 146 Collar 203
Capsule wall 178 Cololejeunea 117
Carbohydrates 284 Columella 117, 151, 322
Carbon and nitrogen cycling 304 Coma 112
Carbon cycling 304 Community 341
Carbon fixation by bryophytes 304 Comparative tables 364
Carcinogenic N-nitrosamines 298 Comparison of external features of gametophyte 365
Carcinoma 285, 355 Comparison of female sex organs 368
Cardiotonic 300 Comparison of male sex organs 367
Carex 150 Comparison of the internal structure 366
Carotenoids 284 Comparison of the sporophyte 369
Carpocephalum 103 Condensation 112
Carrpos 82 Condensation theory 7
Categories of dehiscence of capsule 213 Conducting parenchyma 324
Categories of hormones 229 Conducting strand 320, 323
Caulid 143 Conduction in bryophytes 276
Cauloid 180 Conduction in sporophytes 321
Caulonema 168 Conduction of water 320
Censor mechanism 216 Conocephalum 7
Central cylinder 168 Copper mosses 286, 290
388 � Index
Corsinia 82 Distichum 283

Corsiniaceae 173 Ditrichium 294
Cover cell 10, 105 Ditrichum 307, 308
Cratoneura 342 Divergent branches 151
Cryphaea 276, 351 Drepanocladus 207
Cryptogams 200 Drepanolejeunea 203
Cryptomitrium 109 Drepanophyllum 276
Cryptopolar 5 Drooping branches 151
Cryptothallus 81 Drummondia 203
Cushions 342 Dryptodon 290
Cyanobacteria 307 Dumortiera 7
Cyathodium 145
Cyathophorum 230 E
Cytogenetics 329
Ecology 340
Cytokinins 282
Ecology of bryophytes 340
Cytokinins 329
Economic importance of bryophytes 353
Cytology 329
Ectohydric 202, 12
D Ectohydric 276
Ectropothecium 201
Dawsonia 5, 197, 199, 203, 219, 284, 285, 347 Egg 157
Day-neutral bryophytes 227 Elater 9
Death and decay of older parts 258 Elaterophore 57, 74
Dehiscence 19 Elaters 57, 208
Dendroceros 18, 194, 322 Elaters 79, 57
Dendroligotrichum 284 Embryo 9, 92, 108
Dermatitis 296 Embryobionta 1
Desmatodon 270, 200, 331 Embryogeny 50, 147
Detrichium 341 Embryonic layers 266
Deuters 325 Embryophyta 1, 275
Development of the capsule 175 Embryophytes 243
Diaphragm 179 Encalypta 215, 194
Dicranella 207 Endohydric 203, 223
Dicranum 200, 359 Endospore 45
Differences between algae and bryophytes 13, 14 Endosporium 94
Differences between bryophytes and pteridophytes 15 Endostome 182
Differences between three classes of bryophyta 18 Endothecial 19
Differentiation stage 211 Endothecium 43, 211
Dioecious 144 Entodon 356, 357
Diphyscium 276 Enzymes 279
Diploid 10 Ephemeropsis 5
Diploid generation 197 Ephemerum 142
Diplolepideae 215 Epiphragm 189
Diplolepidous 146 Equisetum 112
Diplophyllin 285 Erosion control by bryophytes 287
Diplophyllum 201, 258, 354 Ethylene 282
Diplophyllum albicans 211, 355 Eucalypta 210
Disc 100 Euchromatin 332
Dispersal of spores 180 Eurhynchium 5
Dissolved organic carbon 306 Evolution of gametophytes in bryophytes 311
Distichium 200 Evolution of the sporophyte 314
Index � 389
Exine 94 Geothallus 70
Exoscopic 12 Germ disc 94
Exospore 45 Giant-thallose-liverworts 77
Exosporium 93 Gibberellins 282, 230
Exostome 182 Glossary 371
Explosive mechanism 159 Granite moss 163
External conduction 321 Granite mosses 141
Green stage 210
F Grimaldone 295
Grimmia 5
Fate of amphithecium 177
Growth-form 342
Fate of archesporium 266
Growth-form classification 276
Fate of endothecium 177
Growth substances 282
Fatty acid 283
Gymnocoba 290
Fertilization 8, 143, 145, 203, 143 228
Gymnocolea 297
Fimbriaria 209, 141, 321
Gymnomitrion 344
Fissidens 200, 9, 166, 342
Gymnostomous mosses 214
Fissidense 338
Flagelliform branches 151 H
Flavones 283
Flavonoids 283 Habitat 343
Floribundaria 342, 357 Hadrome 153
Folioceros 245 Hair-cap-moss 210
Foliose 5, 45 Hanging bryophytes 276, 277
Fontinalis 5, 199, 202, 357 Hapalosiphon 307
Foot 6, 107, 166, 167 Haplocladium 294, 354
Fossil bryophyta 349 Haploid 10
Fossombronia 7 Haploid generation 197
Fossombroniaceae 52 Haploid spores 12
Four-toothed moss 183 Haplolepideae 215
Fragmentation 7, 38 Haplolepidous 146
Frullania 7, 355 Haplomitrineae 35
Frullaniaceae 36 Haplomitrium 28, 21
Funaria 6 Haplozia 259
Hapteres 223
G Haustorium 42
H-chromosomes 334
Gametophyte 5, 140, 197, 232, 233
Hedwigia 203
Gametophytic generation 10
Helodium 201
Gemma 82, 61, 183, 184, 259, 253
Hepatica 22
Gemma cups 99
Hepaticae 2, 232
Gemmae 7, 47, 259
Hepaticites 349
Gemmae of hornworts 260
Hepaticophyta 3
Gemmae of liverworts (Hepaticopsida) 259
Hepaticopsida 1, 136
Gemmae of mosses 260
Hepatites 350
Gemmae of some bryophytes 260
Hepatophyta 3
Gemmaling 75
Herbertia 199
GeneBank database 4, 244
Herbertus 357
Genetic engineering of bryophytes 359
Heterochromatin 330, 332
Genosystematics 19
Heterochromosome 330
Geobelobryum 210
Heteromorphic 233
Georgia 270
390 � Index
Heteromorphic alternation of generations 10 Irish moss 2

Heteropycnotic 330 Isomorphic 233
Heteropycnotic chromosomes 334 Isomorphic alternation of generations 232
Heterosporous 218 Isothecium 278
Homalothecium 284
Homogeneous 118 J
Homologous 233
Jacket 170
Homologous theory 241
Jamesoniella 307
Homosporous 10
Jubula 114
Horned liverworts 17
Jugermanniaceous gametophytes 34
Horneophyton 138, 291, 277
Juncus 150
Hornworts 1, 19
Jungagia 352
Humus 345
Jungermannia 287
Hyaline cells 152
Jungermanniales 35
Hyalodermis 153
Jungermanniales acrogynae 35
Hybrid 336
Jungermanniales anacrogynae 35, 52
Hybridization 336
Jungermannineae 20
Hydroids 4, 326
Jungermanniopsida 148
Hydrom 190
Jungermannites 351
Hydrom cylinder 192
Hydrom mantle 192
K
Hydrotropism 280
Hygrohypnum 292 Karyogamy 145
Hygroscopic mechanism 213
Hylocomium 199, 288, 355, 356 L
Hymenophyton 283
Hymenophytum 202 Lamellae 190, 320
Hypnum 215, 283, 356 Lantern mosses 162
Laptobryum 227
I Large-sized spores 186
Large-spored mosses 185
Incubous arrangement 36 Lateral scales 74
Incubous type 38 Leaf 169
Indicators of environmental conditions 286, 287, 288 Leaf cladia 258
Influence of temperature on sexuality 228 Leafy forms 199
Initiation of buds 252 Leafy forms of bryopsida 199
Innate 224 Leafy forms of hepaticopsida 198
Inner spore sac 176 Leafy gametophore stage 148
Innovation 151 Leafy liverworts 2
Innovation organs 164 Leafy stage 142
Innovations 7, 154 Leaves 143, 152
Insect antifeedant 297 Lejeunea 200, 276, 310
Intercalary meristem 127 Lepidopilum 203
Intercalation theory 234 Lepidozia 199, 310, 355
Internal conduction 322, 327 Leptobryum 143, 228, 230
Interpolation theory 234 Leptodictyum 291
Interrelationships 248 Leptoid mantle 187
Intia 351 Leptoids 4, 324, 325, 326, 327
Intine 94 Leptolejeunea 7
In vivo 270 Leptom 190, 324
Involucre 72 Leptom mantle 192
Index � 391
Leucobryum 5, 321, 356, 358 Mature sporophyte 50, 158

Leucodon 343 M-bivalent 330
Leucolejeunea 221 Medicinal uses of bryophytes 353
Leucoloma 276 Medulla 153
Lichens 16 Megaceros 117
Lieosporoceros 246 Meiospores 9
Life cycle 10, 110 Merceya 290
Life-cycle in bryophytes 11 Meristematic zone 127
Life cycle of Riccia 95 Mesosporium 94
Life history of Anthoceros 130 Metabolism 254
Lignin 3, 283 Metzgeria 7
Ligulate scales 96 Metzgeriaceae 52
Lipids 282 Metzgerineae 35
Liverwort 1, 2, 19, 254, 285 Micro-chromosomes 334
Living fossil in hepaticae 28 Microdus 230
Long-day bryophyte 226 Microlepidozia 199
Lophocolea 213, 295 Micromitrium 207
Lophocolea cuspidata 211 Microscopic moss 181
Lophozia 259 Midribs 325
Lunularia 7, 344 Mielichhoferia 287
Lunularic acid 282 Mnium 200, 353
Lycopodium 2 Mnium medium 144
Modification theory 241
M Molecular phylogenetics 248
Molecular studies 245
Macrocystis 348
Monocarpus 5
Macromitrium 218, 7, 9
Monoclea 24
Macrothamnium 356
Monocleaceae 52
Madotheca 36
Monocleales 77
Madothecaceae 36
Monoecious 144
Makednothallus 202
Monophyletic 346
Mannia 81, 22
Monoselenium 311
Marchantia 7, 12, 16, 17, 81, 198, 353, 354
Morphogenesis 250
Marchantiaceae 82, 55
Moss 140
Marchantiaceous gametophytes 114
Moss bag 288
Marchantiales 77, 2, 89, 103, 123, 155, 244, 246
Mosses 1, 19
Marchantia polymorpha 211
Mosses and liverworts of medicinal importance 293
Marchantin-A 297, 300
Moss gametophore 143
Marchantiolites 351
Moss gardens 359
Marchantiophyta 3, 16, 136
Mosslike hepatics 28
Marchantiopsida 20, 135, 77
Moss plant 142
Marchantites 351
Moss temple 359
Marsupella 297
Movement 279
Marsupium 77, 206, 207,
Mucilage hairs 26
Mastigophora 298
Mucilaginous hairs 201
Mastigophorene 298, 299
Multiform-thallose-hepatics 52
Mastigophorene-A 300
Musci 2
Mats 342
Muscites 351
Mature antheridium 49
Muscle-relaxing activity 299
Mature archegonium 49, 174
Mylia 207
Mature gametophyte 197
Myxohydric 275, 276
Mature sporogonium 44, 63
392 � Index
N Orthodontous peristome 146

Orthotrichum 213, 315, 334
Naiadita 312, 350, 351 Osteoporosis and allergy 299
Nanobryum 223 Ostiole 101
Nanomitrium 203 Oxymitra 83
Nardia 220, 344
Neck 105 P
Neckera 203, 343, 356, 359
Nematodontous peristome 146 Palaeohypnum 352
Neohattoria 45 Pallavicinia 202, 18, 172, 188, 192, 52, 354, 355
Neohodgsonia 219 Pallavicinites 350
Neuroloma 163 Pallaviniaceae 52
Neurotrophic properties 298 Panonychus 297
N fixation 307 Papillae 201
Nitella 349 Paraphyllia 201, 320
Nitrogen-fixing organisms 288 Paraphyly of bryophytes 244
Nitrogen metabolism in bryophytes 278 Paraphyses 144, 170, 172, 188
Non-sexual generation 10 Paroicous mosses 144
Nontracheophytes 3 Passive dehiscence and passive dispersal 213
Norwallia 211 Peat 150
Nostoc 118 Peatcork 357
Nostoc - cavities 120 Peatcrete 357
Nostoc colonies 132 Peatfoam 357
Notothylaceae 117 Peat moss 113, 141, 151
Notothylas 19 Peatwood 357
NPP 305 Pellia 3
Nucleolar chromosomes 336 Pelliaceae 52
Number of spores 219 Peltolepis 202, 143
Number of spores per plant 211 Penicillium 297
Nurse cells 74 Perianth 48
Perichaetial leaves 160
O Perichaetium 104
Perigonial bracts 48
Oblique septa 166, 167, 168 Perigonium 181
Octoblepharum 251, 262, 276, 358 Perigynium 104, 228
Oedipodiopsida 20, 246 Perinium 45
Oedogonium 234, 235 Perispore 93
Oil cells 99 Peristome 117, 127, 128, 137, 146, 179
Oligotrichum 216 Peristome architecture 247
Omphalanthus 202 Peristomial teeth 148, 210, 214
Oogamous 7 Pesticidal properties 298
Oospore 106 Petalophyllum 65
Operculum 145, 176 pH 229
Organic acids 279 Phaeoceros 117, 189, 135, 137
Organic acids in bryophytes 284 Phascum 203, 189
Organic compounds from bryophytes 281 Philonotis 211, 306, 142, 353, 357
Origin of anthocerotae 138 Photoperiod 226
Origin of bryophytes 346 Photoperiodism 226
Origin of bryophytes from algae 349 Photosynthesis in bryophytes 277
Origin of elaters 268 Photosynthetic filaments 97
Origin of land flora 348 Photosynthetic region 83
Origin of pteridophytes 135
Index � 393
Phragmoplast 349 Products of archesporium 264

Phyllids 6, 223, 331 Progressive death and decay 87
Phyllogonium 305 Progressive death and decay process 121
Phylloid 5, 6 Progressive evolution 111, 351
Phylloid cladia 7 Progressive simplification 318
Phylogenetic tree of liverworts 246 Protonema 10
Phylogenetic trees 244, 349 Protonemal bryophytes 277
Physcomitrella 251 Protosphagnum 351
Physcomitrium 210, 335 Pseudobarbella 342
Physiology of bryophytes 275 Pseudoelater mother cells 127
Phytochrome 254 Pseudoelaters 117
Piliferous layer 190 Pseudoparianth 104, 106
Pin-cushion moss 321 Pseudopodium 150, 159, 347
Plagiochasma 81 Pseudoscleropodium 342
Plagiochila 199, 246, 354, 357 Psilophytalean ancestry of anthocerotales 138
Plagiomnium 227, 344 Psilophytales 138
Plagiopodopsis 352 Psilopilum 287
Plagiopus 294 Psilotum 241
Plagiothecium 201 Pteridophyta 135
Plant body 5 Pteridophytean origin of marchantiales 112
Plasmogamy 145 Pteridophytes 1, 13
Platyhyphidium 284, 291 Pteridophytic origin 347
Pleuridium 211 Pterygynandrum 260
Pleurocarpous 140 Ptilidium 344
Pleurocarpous gametophytes 140 Ptychantus 300
Pleurocarpous mosses 142, 200 Ptychomitrium 338
Pleurozium 287 Pungent and bitter substance 295, 296
Podomitrium 202 Pyrenoid 13, 116
Pogonatum 186, 347
Pohlia 143 R
Poikilohydry 302
Radial strands 190
Polarity 224
Radula 7, 199, 211, 212, 221, 276, 283, 285, 296,
Polyphyletic origin of bryophytes 346, 311
297, 300, 311
Polyploids 337
Rays 104
Polyploidy 337
Reboulia 7, 202, 219, 220, 228, 230, 258, 268, 281,
Polyssaievia 351
300, 322, 354
Polytrichaceae 186
Rebouliaceae 82
Polytrichales 186
Recapitulation 349
Polytrichiidae 186
Receptacle 78, 114
Polytrichium 258
Reduction 112
Polytrichopsida 20
Reduction theory 112, 59
Polytrichum 5
Regeneration 11, 257, 261
Porella 9
Regressive evolution 240
Porellaceae 36
Reindeer moss 2
Postical lobe 37
Resemblances between liverworts and mosses 147
Pottia 214
Resemblances of Sphagnum 161, 70
Prehepatics 348
Respiration in bryophytes 278
Preissia 81
Respiratory quotient 278
Primary archegonium 155
Retort cells 153
Primary pit fields 322
Retrogressive evolution 111, 167
Primary protonema 7, 143
394 � Index
Rhacocarpus 3 Secondary protonema 7, 143, 138

Rhacomitrium 276, 83 Seed beds 359
Rhacopilum 356 Selaginella 244
Rhizogonium 351 Sematophyllum 276
Rhizoid 4, 96, 200 Senescence 255
Rhizoidal branches 147 Senescence hormone 282
Rhizoidal cells 99 Seta 6, 9, 108, 109
Rhizoid production 255 Sewardiella 65
Rhizome 27 Sex chromosomes 334, 335
Rhizomnium 359 Sex determination 253
Rhocopilum 199 Sex expression 252
Rhodobryum 262, 354 Sexuality 226
Rhyncostegium 284 Sexual reproduction 7
Rhynia 138 Short-day bryophytes 227
Rhytidiadelphus 342, 356 Simodon 65
Rhytidiopsis 358 Size of the spores 219
Riccardia 7 Smooth-walled rhizoids 84, 85, 97
Riccardiaceae 52 Spanish moss 2
Riccardin-C 300 Special bivalent 330
Riccardius 60 Spermatogenesis 90, 102
Riccia 5, 309, 353 Sphaerocarpaceae 70, 250
Ricciaceae 82 Sphaerocarpales 70
Ricciocarpos 5 Sphaerocarpos 70
Ricciopsis 351 Sphaerosporoceros 245
Riella 5 Sphagnales 141
Riellaceae 70 Sphagnophyta 19
Rim 179 Sphagnopsida 20
Role of bryophytes in C and N cycling 309 sphagnorubins 283
Roof 121 Sphagnum 5, 354, 355, 357
Root 4, 27 Spiral-ring mechanism 213
Rubisco 307 Spiridens 277
Rudula 260 Splachnum 142, 219, 220
Spodoptera 297
S Sporangium 205
Spore 45, 93
Sacculatal 297
Spore germination 109, 129
Salairia 351
Sporeling 220
Salient features of bryophytes 11
Spore mother cells 127
Sargassum 348
Spore sac 177
Sauteriaceae 82
Spore viability 224
Scales 81, 85, 154, 160, 169, 307
Sporocytes 93
Scapania 199, 7, 145
Sporogenesis 93
Scapania undulata 211
Sporogon factor 273
Scents 295
Sporogonium 9
Schiffneria 311
Sporogonium in Anthoceros 126
Schistochila 207
Sporogonium of Riccia and Marchantia 111
Schistostega 223, 74
Sporophyte 6, 158
Schlotheimia 218
Sporophyte in bryophytes 205, 237
Scopelophila 287
Sporophyte of Anthoceros 238
Scouleria 215
Sporophytic generation 10
Secondary archegonia 155
Stalk 100
Index � 395
Stem 143 Tortula 216, 245, 338, 344

Stem cladia 258 Torula 5, 207
Stephensoniella 200 Totella 343
Stereids 183 Trabeculae 178
Stereom 190 Tracheophyta 1
Stereome 324 Trachycystis 359
Steroids 283 Trachypodopsis 356, 357
Stictolejeunea 221 Transfer cells 210
Stolon 27 Transformation theory 241
Stomata 127 Treatment of Sarcoma-37 285
Storage region 86 Treubia 198
Stylus 46 Treubiaceae 52
Succession 343 Treubiopsida 20
Suspensor 42 Trichocolea 201
Symphyogyna 327 Trichocoleopsis 296
Synecology 341 Trichophyton 297
Synoicous mosses 144 True liverworts 17
True mosses 141
T Tuber 259
Takakia 24, 25, 26, 28, 198, 199, 200, 201, 203, 295, Tuberculate rhizoid 85, 97
322, 323, 327 Tuber-forming species 259
Takakiaceae 25 Tubers 7, 88
Takakiales 24, 25 Tuft-forming 276
Takakiopsida 20, 246 Turf moss 151
Tamariscol 295 Turfs 342
Targionaceae 82, 202 Tympanum 189
Targionia 81, 7, 295, 314, 316, 319, 337
Terpenoids 283, 293
U
Tetraphidaceae 183 Ulota 203, 260, 276, 343
Tetraphidales 183 Ulothrix 28
Tetraphidopsida 20, 198 Unique characters of Sphagnum 162
Tetraphis 183 Uskatia 351
Tetraplodon 336
Thalloid 5, 198 V
Thalloid bryophytes 198
Thalloid hepatics 82 Vaginula 159, 263
Thalloid liverworts 2, 175 Vascular elements 134
Thallophyta 1 Vasopressin antagonist activity 300
Thallophytes 1 Vegetative propagation 183, 253
Thallus 131, 198 Vegetative reproduction 6, 87, 257
Thamnium 283, 342 Venter 157
Theca 117, 145, 146 Venter canal 157
Theory of origin of strobilus 134 Ventral scales 74
Theory of progressive evolution 113 Vessels 4
Theory of progressive sterilisation 318 Viridivellus 5
Theory of sterilization 314 Vorcutannularia 352
Thuidium 201, 342
Tillandsia 2 W
Timmiella 281, 354, 355 Wardia 215
Tmesipteris 241 Water relations 275
Tortella 222, 345 Water-rupture mechanism 213
396 � Index
Wefts 342 Young gametophytic thallus 109

Weisnerella 311 Young sporophyte 50, 51
Weissia 336, 344
Z
Y
Zoopsis 197, 203, 311
Young gametophore 180, 181 Zygodon 343
Young gametophyte 65, 160 Zygote 8

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Series on Diversity of Microbes and Cryptogams

McGraw Hill Education (India) Private Limited

McGraw Hill Education Offices

Series on Diversity of Microbes and Cryptogams: BryOPhyta

1.1 What are Bryophytes? 1

4.1 General Characteristics 34

5.1 Sphaerocarpales and their General Characteristics 70

6.1 What are Monocleales? 77

7.1 Limits of Marchantiales? 81

8.1 What is Anthoceropsida? 116

9.1 What is Bryopsida? 140

10.1 Sphagnales or Bog Mosses 150

11.1 What is a Gametophyte? 197

12.1 What is a Sporophyte and a Spore? 205

13.1 What is Technically a Spore? 218

14.1 Major Factors Affecting Sexuality 226

15.1 What is Alternation of Generations? 232

16.1 Current Status of Molecular Studies on Bryophyte Phylogeny 243

17.1 What is Morphogenesis? 250

18.1 Difference between Vegetative Reproduction and Regeneration 257

21.1 Water Relations 275

22.1 Antibiotics 281

24.1 What are Biologically Active Compounds? 293

25.1 Role of Plants in Regulating Biogeochemical Cycles 302

26.1 Views on the Evolution of the Gametophyte in Bryophytes 310

27.1 What is Evolution of the Sporophyte in Bryophytes? 314

28.1 How do Bryophytes Maintain Poor Internal Conducting Strands? 320

29.1 Some Basic Techniques of Cytogenetics 329

31.1 Origin of Bryophytes 346

32.1 Utility of Bryophytes: A General Overview 353

The present book on Bryophyta includes 32 chapters.

Some major highlights of the book are as follows:

WHAT ARE BRYOPHYTES? 1.1

1.1.1 Are all Mosses Bryophytes?

1.1.2 What are Liverworts?

HOW DO RECENT SYSTEMATISTS TREAT THE TERMS

WHY ARE BRYOPHYTES ALWAYS SMALL SIZED

WHY DO BRYOPHYTES POSSESS RHIZOIDS

NUMBER OF SPECIES AND GENE BANK DATABASE 1.5

AGE OF BRYOPHYTES 1.6

OCCURRENCE AND DISTRIBUTION 1.7

GAMETOPHYTE PLANT BODY 1.8

VEGETATIVE REPRODUCTION 1.9

SEXUAL REPRODUCTION 1.10

Fig. 1.4 An archegonium of Riccia just before fer liza on

Fig. 1.5 A mature sporophyte of Porella

YOUNG GAMETOPHYTE 1.12

LIFE CYCLE ALTERNATION OF GENERATIONS 1.13

Fig 1.6 Diagramma c representa on of the life cycle in bryophytes

A NOTE ON REGENERATION IN BRYOPHYTES 1.14

SALIENT FEATURES OF BRYOPHYTES: AT A GLANCE 1.15

3. Their plant body is gametophytic, which is independent.

AFFINITIES OF BRYOPHYTES 1.16

1.16.1 Aﬃni es of Bryophytes with Algae

2. Diﬀerences between Bryophytes and Algae

1.16.2 Aﬃni es of Bryophytes with Pteridophytes

Table 1.1 Diﬀerences between algae and bryophytes

S. NO. ALGAE BRYOPHYTES

2. Diﬀerences between Bryophytes and Pteridophytes

Table 1.2 Diﬀerences between bryophytes and pteridophytes

S. NO. BRYOPHYTES PTERIDOPHYTES

ORIGIN OF BRYOPHYTES 1.17

TEST YOUR UNDERSTANDING

1. Who was the ﬁrst to introduce the word “bryophyta”?