Bryophyta General Characters

• Bryophytes (nonvascular plants) are the only embryophytes

(plants that produce an embryo) whose life history includes a
dominant gametophyte (haploid) stage.
• Bryophyta (Gr. Bryon = moss; phyton = plant), a division of
kingdom Plantae comprises of Mosses, Hornworts and
Liverworts.
• These plants occupy a position between the thallophytes (Algae)
and the vascular cryptogams (Pteridophytes).
• Bryophytes produce embryos but lack seeds and vascular
tissues.
• They are the most simple and primitive group of Embryophyta.
• Bryophytes grow in two habitats i.e. water and land so known as
amphibians of plant kingdom.
• They are said to be the first land plants or non-vascular land
plants.
• They cannot reproduce without sufficient moisture because
without water sex organs neither matures nor dehisces.
• Presence of swimming antherozoids is an evidence of their
aquatic ancestory.
• They comprise three main taxonomic groups:
• mosses (Bryophyta), liverworts (Marchantiophyta or
Hepatophyta) and hornworts (Anthocerotophyta)
• Most bryophytes have erect or creeping stems and tiny leaves,
but hornworts and some liverworts have only a flat thallus and no
leaves.
• There are possibly 25,000 species of mosses and liverworts all
over the world.
Origin of Bryophytes
• Nothing definite is known about the origin of Bryophytes because
of the very little fossil record.
• There are two views regarding the origin of Bryophytes.
(i) Algal Hypothesis of Origin:
• There is no fossil evidence of origin of Bryophytes from algae but
Bryophytes resemble with algae in characters like-amphibious
nature, presence of flagellated antherozoids and necessity of
water for fertilization.
• Photosynthetic pigments like Chl a, b, carotene, leutine and
violaxanthine.
• Starch as reserve food material.
• Filamentous protonema in the juvinile stage.
• This hypothesis was supported by Bower (1908), Fritsch (1945)
and Smith (1955) etc.
• According to Fritsch (1945) and Smith (1955) Bryophytes have
been originated from the heterotrichous green algae belonging to
the order Chaetophorales for e.g., Fritschiella, Coleochaete and
Draparnaldiopsis.
ii) Pteridophytean Hypothesis of Origin:
•According to this hypothesis Bryophytes are descendent of
Pteridophytes.
•They are evolved from Pteridophytes by progressive simplification
or reduction.
•This hypothesis is based on certain characters like-presence of type
of stomata on the sporogonium of Anthoceros and apophysis of
mosses similar to the vascular land plants, similarly in the
sporophytes of some Bryophytes (e.g., Anthoceros, Sphagnum,
Andreaea) with some members of Psilophytales of Pteridophytes
(e.g., Rhynia, Hormophyton etc.).
•Similarity in photosynthetic pigments.
•Cell wall structure.
•Food reserves.
•Reproduction methods.
•This hypothesis was supported by Kashyap (1919), Christensen
(1954), Proskaner (1961), Mehra (1969) etc.
GENERAL CHARACTERISTICS
1. Small group of primitive land dwellers having small leafy or
thalloid green plant body.
2. Like thallophytes plant body is gametophytic, independent,
dominant, autotrophic, either thalloid (i.e., thallus like, not
differentiated into root, stem and leaves) or foliose, containing a
rootless leafy shoot.
3. Plant body is very small and ranges from a few mm too many cm.
Eg: Zoopsis is the smallest bryophyte (5 mm.) while the tallest
bryophyte is Dawsonia (50-70 cm.).
4. Leaves and stems like structures as ‘axis’ and ‘phylloid’
respectively.
5. Roots are absent.
6. Functions of the roots are performed by rhizoids.
7. Cells are also capable to absorb moisture directly from the ground
or atmosphere.
8. Rhizoids may be unicellular, un-branched (e.g., Riccia,
Marchantia, Anthoceros) or multicellular and branched (e.g.,
Sphagnum, Funaria).
9. In members of order Marchantiales (e.g., Riccia, Marchantia)
scales are present. These are violet coloured, multicellular and
single cell thick. They protect the growing point and help to
retain the moisture.
10. Vascular tissue (xylem and phloem) is completely absent.
11. However, in some Bryophytes (e.g., mosses) a few cells in groups
of 2-3 are present for conduction of water and translocation of
food (photo assimilate).
12. These cells are known as hydroid (collectively hydrom) and
leptoids respectively.
13. Cuticle and stomata are absent.
Distribution
• Bryophytes are distributed throughout the world, from polar and
alpine regions to the tropics.
• Bryophytes do not live in extremely arid sites or in seawater,
although some are found in perennially damp environments
within arid regions and a few are found on seashores above the
intertidal zone.
• A few bryophytes are aquatic.
• Bryophytes are most abundant in climates that are constantly
humid and equable.
• The greatest diversity is at tropical and subtropical latitudes.
• Bryophytes dominate the vegetation of peat land in extensive
areas of the cooler parts of the Northern Hemisphere.
• The geographic distribution patterns of bryophytes are almost
cosmopolitan.
• Bryophytes are represented by 960 genera and 25,000 species.
• In India, Bryophytes are quite abundant in both Nilgiri hills and
Himalayas; Kullu, Manali, Shimla, Darjeeling, Dalhousie and
Uttarakhand are some of the hilly regions which also have a
luxuriant growth of Bryophytes.
• Eastern Himalayas have the richest in bryophytic flora.
• A few species of Riccia, Marchantia and Funaria occur in the
plains of U.P., M.P. Rajasthan, Gujarat and South India.
• In hills they grow during the summer or rainy season. Winter is
the rest period.
• In the plains the rest period is summer, whereas active growth
takes place during the winter and the rainy season.
• Some Bryophytes have also been recorded from different
geological eras e.g., Muscites yallourensis (Coenozoic era),
Intiavermicularies, Marchantia spp. (Palaeozoic era) etc.
HABITAT
• Bryophytes grow densely in moist and shady places and form
thick carpets or mats on damp soils, rocks, bark of trees especially
during rainy season.
• Majority of the species are terrestrial but a few species grow in
fresh water (aquatic) e.g., Riccia fluitans, Ricciocarpos natans,
Riella etc.
• Bryophytes are not found in sea but some mosses are found
growing in the crevices of rocks and are being regularly bathed by
sea water e.g., Grimmia maritime.
• Some bryophytes also grow in diverse habitats e.g., Sphagnum-
grows in bogs, Dendroceros-epiphytic, Radulaprotensa-
epiphyllous, Polytrichum juniperinum- xerophytic, Tortula muralis-
on old walls.
• Tortula desertorum in deserts, Porella platyphylla-on dry rocks,
Buxbaumia phylla (moss) and Cryptothallus mirabilis (liverwort)
are saprophytic
LIFE CYCLE
• The life cycle of bryophytes shows two distinct phases namely a
haploid gametophytic phase and a diploid sporophytic phase
alternating with each other.
• The adult plant body represents the gametophyte.
• A short-lived sporophyte occurs as a parasite on the
gametophyte.
• The gametophyte is the sexual phase in the life cycle of
bryophytes.
• It develops sex organs that produce gametes.
• In bryophytes (mosses, liverworts, and hornworts), the
gametophyte is the most visible stage of the life cycle.
• The bryophyte gametophyte is longer lived, nutritionally
independent, Sporophytes are typically attached to the
gametophytes and nutritionally dependent on them.
REPRODUCTION IN BRYOPHYTES
• Bryophytes reproduce only by vegetative and sexual means.
• Asexual reproduction is absent in these.
A. Vegetative Reproduction in Bryophytes:
• The vegetative reproduction takes place in favourable season for
vegetative growth.
• Majority of the bryophytes propagate vegetatively and it is
brought about in many ways.
By Death and Decay of the Older Portion of Thallus or by
Fragmentation:
• In Bryophytes the growing point is situated at the tip of the
thallus.
• The basal, posterior or older portion 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.
• These detached lobes or fragments develop into independent
plants by apical growth.
• This is the most common method of vegetative reproduction in
Riccia, Marchantia, Anthoceros and some mosses like
Sphagnum.
By Persistent Apices:
• Due to prolonged dry or summer or towards the end of growing
season the whole thallus in some bryophytes (e.g., Riccia,
Anthoceros, Cyathodium) dries and get destroyed except the
growing point.
• Later, it grows deep into the soil and becomes thick.
• Under favourable conditions it develops into a new thallus.
By Regeneration:
• The liverworts possess an amazing power of regeneration.
• Part of the plant or any living cell of the thallus (e.g., rhizoid,
scales) are capable of regenerating the entire plant. e.g., Riccia,
Marchantia etc.
By Tubers:
• Tubers are formed in those species which are exposed to desiccations
(drying effect of the air).
• Towards the end of the growing season, the subterranean branches get
swollen at their tips to from the underground tubers.
• On the periphery of a tuber are two to three layers of water proof, corky,
hyaline cells develop.
• These layers surround the inner cells which contain starch, oil globules and
albuminous layers.
• During the unfavorable conditions the thallus dies out but the dormant
tubers remain unaffected.
• On the return of the favourable conditions each tuber germinates to form a
new plant e.g., Riccia, Anthoceros, Fossombronia etc.
By Gemmae:
• Gemmae are multicellular
reproductive bodies of various
shapes and colours.
• These are produced in gemma
cups, on the surface and axil of
the leaves, on stem apex or even
inside the cells.
• They get detached from the
parent plant and after falling on
a suitable substratum gemmae
give rise to a new individual
directly (e.g., Marchantia) or
indirectly (e.g., Mosses).
By Adventitious Branches:
• The adventitious branches
develop from the ventral
surface the thallus e.g., Riccia
fluitans and Anthoceros.
• On being detached from the
parent plant these branches
develop into new thalli.
• In Marchantia, Dumortiera
these branches develop from
archegoniophore while in Pellia
these branches arise from the
dorsal surface or margins of the
thallus called automnal fronds.
By Innovation:
• In Sphagnum one of the branches in the apical cluster instead of
forming drooping branches or divergent branches, develop more
vigorously than the others and continues the growth upwards.
• This long upright branch has all the characteristics of main axis. It
is called innovation.
• Due to progressive death and decay of the parent plant these
innovation become separated from the parent plant and establish
themselves as parent plants.
By Rhizoidal Tips:
• The apical part of the young rhizoids divide and re-divide to form
a gemma like mass of cells e.g., Riccia glauca.
• These cells contain chloroplast and are capable to develop into
new thallus.
8. By Primary Protonema:
• Primary protonema is the filament like stage produced by the
developing spores of the mosses.
• It produces the leafy gametophores.
• It breaks into short filament of cells by the death of cells at
intervals.
• Each detached fragment grows into a new protonema which
bears a crown of leafy gametophores e.g., Funaria.
9. By Secondary Protonema:
• The protonema formed by other methods than from the
germination of spores is called secondary protonema.
• It may develop from any living cells of the leafy gametophore
i.e., from leaf, stem, rhizome, injured portion of the leafy
gametophore, antheridium, paraphysis or archegonium.
• From this arise the leafy gametophores or lateral buds in the
same manner as in primary protonema e.g., Funaria, Sphagnum.
10. By Bulbils:
• These are small resting buds develop on rhizoids.
• Bulbils are devoid of chlorophyll but full of starch.
• On germination bulbils produce a protonema which bears leafy
gametophores
11. By Apospory:
• The production of diploid gametophyte from the unspecialized
sporophyte without meiosis is known as apospory e.g.,
Anthoceros.
• In Funaria green protonemal filaments bear lateral buds which
develop into leafy gametophores
B. Sexual Reproduction in Bryophytes
• Sexual reproduction is highly oogamous.
• Male and female sex organs are known as antheridia (Sing,
antheridium) and archegonia (Sing, archegonium), respectively.
• Sex organs are jacketed and multilayered.
• Antheridium is stalked, pear shaped or oblong and has an outer
one cell thick jacket which encloses a mass of fertile cells called
androcytes.
• Each androcyte metamorphoses into biflagellate antherozoid.
• Archegonium is stalked, flask shaped structure.
• It has a basal swollen portion called venter and an elongated
neck.
• The neck is filled with many neck canal cells whereas venter has
a large egg cell and a small venter canal cell.
• Antherozoids are attracted towards the neck of the archegonium
chemotactically by certain substances (like sugars, malic acid,
proteins, inorganic salts of potassium etc.) present in the
mucilaginous substance formed by the degeneration of neck
canal cells and venter canal cell.
• Water is essential for fertilization.
• The fertilized egg or zygote is the beginning of the sporophytic
phase.
• It is retained within the venter of the archegonium.
Sporophyte
• Without resting period, the zygote undergoes repeated divisions
to form a multicellular structure called the embryo.
• The first division of the zygote is always transverse and the outer
cell develops into embryo. Such an embryogeny is called
exoscopic.
• Embryo develops into a sporophyte or sporogonium.
• The sporophyte is usually differentiated into foot, seta and
capsule.
• In certain cases it is represented only by capsule (e.g., Riccia) or
by foot and capsule (e.g., Corsinia).
• Sporophyte is attached to parent gametophytic plant body
throughout its life.
• It partially or completely depends on it for nutrition.
• Foot is basal, bulbous structure.
• It is embedded in the tissue of parent gametophyte.
• Its main function is to absorb the food material from the parent
gametophyte.
• Seta is present between the foot and capsule.
• It elongates and pushes the capsule through protective layers.
• It also conducts the food to the capsule absorbed by foot.
• Capsule is the terminal part of the sporogonium and its function
is to produce spores.
• All Bryophytes are homosporous i.e., all spores are similar in
shape, size and structure
• Capsule produces sporogenous tissue which develops entirely
into spore mother cells (e.g., Riccia) or differentiated into spore
mother cells and elater mother cells (e.g., Marchantia and,
Anthoceros).
• Spore mother cells divide diagonally to produce asexually four
haploid spores which are arranged in tetrahedral tetrads.
• Elater mother cells develop into elaters (e.g., Marchantia) or
pseudo elaters (e.g., Anthoceros) which are hygroscopic in
nature.
• Elaters are present in liverworts and absent in mosses.
• Venter wall enlarges with the developing sporogonium and forms
a protective multicellular layer called calyptra (gametophytic
tissue enclosing the sporophyte).
Young Gametophyte
1. The meiospore (spore formed after meiosis) is the first cell of the
gametophytic phase.
2. Each spore is unicellular, haploid and germinates into young
gametophytic plant (e.g., Riccia or Marchantia) or first
germinates into a filamentous protonema on which buds are
produced to give rise to a young gametophytic plant, (e.g.,
Funaria).
Alternation of Generation:
• The life-cycle of a bryophyte shows regular alternation of
gametophytic and sporophytic generations.
• This process of alternation of generations was demonstrated for
the first time in 1851 by Hofmeister.
• Thereafter in 1894 Strassburger could actually show the periodic
doubling and halving of the number of chromosomes during the
lifecycle.
• The haploid phase (n) is the gametophyte or sexual generation. It
bears the sexual reproductive organs which produce gametes, i.e.,
antherozoids and eggs.
• With the result of gametic union a zygote is formed which
develops into a sporophyte. This is the diploid phase (2n).
• The sporophyte produces spores which always germinate to form
gametophytes.
Resemblance with Algae
1. Thalloid plant body.
2. Absence of roots.
3. Absence of complex vascular bundles.
4. Autotrophic mode of nutrition.
5. Need of water during fertilization.
6. Presence of chlorophyll pigments.
7. Reserve food material is true starch.
8. Cell wall consists of cellulose which surrounds by a pectic layer
containing galacturonic acid.