“Amphibians of the plant kingdom”

Bryophytes
• A group of non-vascular plants commonly called “the amphibians of plant
kingdom”

• The term Bryophyta was used for the first time by Braun (1864)

• Bryon=moss; phyton=plants (Greek)

• Simplest and most primitive plant groups which occupies a position intermediate
between thallophytes (algae) and pteridophytes
• Comprises Mosses, liverworts and Hornworts

• c. 960 genera and 24,000 species

• Common in moist regions of the world

• Primitive land dwellers

 Distribution and Occurrence
• Thrive in places where sufficient moisture is
available
• Usually occur in humid and shady localities
• Seen in diverse conditions
• Few are permanently submerged in water, few
others thrive on dry rock crevices, wet walls,
decaying logs, etc
• Few are epiphytes on tree trunks
Riccia fluitans
Riella
Cyathodium
Plagiochasma
Frullania
Porella
Fossombronia
Cryptothallus mirabilis
Cryptothallus mirabilis
Plagiochasma Reboulia

Riccia fluitans Conocephalum

Anthoceros
Phaeoceros
 General characters
• Bryophytes have evolved a life cycle which
comprises two phases- Gametophyte and
Sporophyte
• Morphologically distinct (hetermorphic)
alternation of generations
• Gametophyte is haploid and represents the
predominant phase of the life cycle
• Sporophyte is short-living and completely
dependent on the gametophyte
 Gametophyte
• Haploid, green coloured, branched,
independent and autotrophic
• In primitive forms, the gametophyte is
prostrate and thalloid without differentiation
in to roots, stems and leaves
• Roots are absent and fixation and absorption
are effected by root-like and filamentous
rhizoids
• Vascular tissues absent
Dumortiera Marchantia

Lunularia Cyathodium
Aerolae
Scales

Rhizoids

The members of order Marchantiales

possess multicellular scales to protect the
growing regions of the thallus and also to
help in water absorption
• Rhizoids are either unicellular and
unbranched (Hepaticopsida &
Anthocerotopsida) or Multicellular and
branched (Bryopsida)

Unicellular unbranched
rhizoids are of two
types
Smooth walled and
Tuberculate or pegged

Absorb minerals and water, fixation

• In higher bryophytes,
plant body is erect
and differentiated in
to stem (axis) and
lateral appendages
(Leaves)
• Rhizoids are
multicellular and
branched
 Internal structure
• In liverworts, plant body is differentiated in
to 2 distinct regions
Dorsal assimilatory regions
Ventral storage regions
• Dorsal regions encloses many air chambers
• Air canals or channels present in Riccia
instead of chambers
• Air chambers are empty or contains
assimilatory filaments
• Assimilatory filaments contains chlorenchyma
• Each chamber opens to outside by a pore
Storage regions-ventral side
• Composed of colorless parenchyma
• Compactly arranged
• Lack chloroplasts but contains starch
• Oil cells also present
• Lowermost layer of this region bear rhizoids
and scales
 Internal structure
• In hornworts, thallus is very simple without
any cellular differentiation
• In Mosses, stem or axis complex cellular
organization
• Shows three different regions
Epidermis
Cortex
Medulla
Riccardia
 Vegetative Reproduction
1. Progressive death and fragmentation (Riccia,
Marchantia, others)
2. Production of adventitious branches from
underside of thallus and its separation to form
new thalli (Targionia, Riccian fluitans)
3. Gemmae formation (Marchantia, Lunularia)
4. Tuber formation (Riccia bilardieri, R.discolor, R.
vesicata, Exomothica sp. and Cythodium sp.)
5. Persistent growing apices
6. By rhizoids
7. Protonema (Mosses)
 Sexual Reproduction
• Sexual reproduction is Oogamous
• Antheridia- Male sex organ
• Archegonia- Female sex organ
• Homothallic or monoecious- when both male
and female sex organs are developed in the
same thallus (Riccia)
• Heterothallic or dioecious- when male and
female sex organs are produced on different
thalli (Marchantia)
• Occur singly and scattered in a separate cavity on the
dorsal side of thallus (Riccia)
• Sometimes tend to aggregate in groups
• Commonly sex organs are restricted to special
cushion like areas on the thallus- Receptacles
• Receptacles may be sessile or stalked (Marchantia)
• Stalked receptacles are called Gametophores
• Antheridiophores -Antheridia bearing stalked
receptacle
• Archegoniophores -Archegonia bearing stalked
receptacle
In Sphaerocarpales, each antheridium and
archegonium is enclosed in a spherical
covering known as involucre or bottle.
Thus they are known as Bottle liverworts
• In species like Porella, antheridia
are borne on specialized
branches in the axils of leaves
• In Bryopsida, antheridia are
borne in groups, surrounded by
a rosette of leaves called
Perichaetium or perichaetial
leaves which are arranged more
compactly on the axis than the
vegetative leaves
• A large number of multicellular,
capitate hairs are intermingled
with the antheridia, called the
paraphysis
In Bryopsida, antheridia are borne in groups,
surrounded by a rosette of leaves called
Perichaetium or perichaetial leaves which are
arranged more compactly on the axis than the
vegetative leaves

Paraphysis contain chloroplats and protects

from dessication
• Antheridia are globose or elliptical,
stalked
• Consists of a outer one-cell thick
sterile jacket
• Jacket surrounds a central mass of
androcyte mother cells
• Androcyte mother cells divides to
form androcytes
• Androcytes metamorphose in to
antherozoids
• Antherozoids are motile, biflagellate
and usually coiled
• At the tip of the jacket layer there is
one or two relatively large hyaline
cell which forms operculum
 Archegonium
• Flask shaped with a short stalk
• 2 parts- Neck and Venter
• Venter -Basal, swollen portion,
encloses an egg and a venter
canal cell-with a single layered
sterile jacket
• Neck- consists of 5 or 6
longitudinal rows of neck cells
• Neck canal encloses an axial
row of about 6-9 neck canal
cells
• Mouth of the neck is closed by a
rosette of 4 lid or cover cells
Lunularia cruciata
• Archegonia are developed on the dorsal surface
in liverworts and hornworts
• They occur singly or in groups embedded in the
thallus
• In higher groups like Jungermanniales.
Metzgeriales, Sphagnales etc, they occur in
groups
• In Bryopsida, occur as groups in specialized
branches (Archegonial branches)
• They are surrounded by a cluster of leaves called
Perichaetial leaves
• Paraphyses alse present intermingled with
Archegonia
 Fertilization
• Water is essential for fertilization
• The apical cell of the antheridium absorbs water
and bursts, releasing motile antherozoids in a
vesicle
• Vesicle is dissolved in water setting them free
• At the same time neck canal cell and venter canal
cell degenerates forming a mucilaginous mass
• Antherozoids are attracted to the archegonium by
the chemotactic influence of the mucilage
• A large number of antherozoids enter the venter
but only one fuses with egg
 Sporogonium
• Sporophytic or diploid phase of the life cycle
• Simple in structure and radial in symmetry
• Completely dependent on gametophyte
(organically attached)
 Sporogonium
Concerned with the production and
dispersal of haploid spores
(Meiospores)
Consists of 3 parts- Foot, Seta and
Capsule
• Simplest type is seen
in Riccia.
• Represented by a
capsule only
• Foot and seta being
absent
• Sporogonium in Corsinia
is differentiated in to a
basal portion (absorbing
organ) and the capsule
region-Seta is absent
• Sporogenous tissue is
differentiated in to two
types of cells-
• Spore mother cells
• Sterile cells
• Sporogonium in higher taxa has
a slender stalk like structure
called seta in between foot and
the capsule
– Seta remains short until the
spores become mature
– Elongates after maturity thus
forcing the capsule through the
calyptra and placed in a position
favourable for the dispersal of
spores by wind.
– Some of the sporogenous tissue
becomes elaters
Capsule- Spherical or
globose, 1, 2 or more
layered thick capsule wall
Elongated and horn like in
Anthocerotales
Spores + sterile cells
Sterile cells
metamorphose in to
Elaters or pseudoelaters
In higher taxa, a mature
capsule has free spores
inter mixed with elaters
In Anthoceros,
sporophyte is a thin
elongate structure present
on the dorsal surface of
the thallus

Mature sporophyte has

three parts
A foot, capsule and an
intercalary meristematic
zone
• In Bryopsida, the
capsule has a
characteristc
operculum and
peristomial teeth to
help dispersal of spores
• In Anthocerotales and
Bryales, A sterile mass
of tissue is present at
the center of the
capsule called
Columella
• Elaters are long,
slender, spindle
shaped cells with 2 or
3 or sometimes up to
6 bands of spiral
thickening on their
walls
• Elaters are hygroscopic
and organs for spore
dispersal
 Dehiscence
• Sporophyte is partially
autotrophic
• Young sporophyte remain
enclosed within calyptra-a
gametophytic covering
produced from venter of
the fertilized archegonium
• Seta elongate in length as
the capsule matures,
rupturing the calyptra
• Ripe capsule is exposed to
the drying effect of the
surrounding air
• Hygroscopic movements
of thickened cells of the
wall splits the capsule
open by four valves
• The four valves bend back
exposing the spores
 Changes taking place in gametophyte tissue

• Stalk of the archegoniophore elongates

considerably
• Venter cells divide by periclinal walls to form a
2-3 layered Calyptra
• Perigynium forms a cylindrical sheath
• Two lipped curtain like Perichaetium or
involucre is formed
 Nutrition of sporogonium

• Lacks capacity for self nutrition in Riccia;

completely dependent on the parent
gametophyte
• In Marchantia, cells develop chlorophyll during
the differentiation of the young sporophyte.
• Partially dependent during initial stages
• Cells loses chlorophyll upon maturity.
• A mature sporophyte is completely dependent on
gametophye
• Spores are small and
ranges between 10-
30 µm in diameter
• Exine surface is
smooth, papillose or
granulose
• Spores are non-motile and disseminated by
wind
• Each spore germinates in to a young
gametophyte under favourable conditions or
forms a protonema (mosses)
• A protonema is filamentous structure developed
from a spore
• Protonema forms two types of branches
Chloronemal branches
Rhizoidal branches
• Chloronemal branches give rise to many minute
buds which develop to erect gametophytes
 Alternation of generations

• Life cycle of bryophytes is characterised by a

regular alternation of two morpholgically
different gametophytic and sporophytic phases
• Gametophyte is haploid, independent,
autotrophic and bears gametes (thus
gametophyte)
• It develops from the haploid spores produced by
sporophyte
• Gametes represent the last stage of the
gametophytic phase
 Sporophytic phase

• Male and female gametes fuse to form diploid

zygote which functions as the mother cell of
sporophytic phase
• Represented by a sporogonium which
completely depends upon the gametophyte
• Its sporogenous cells undergo reduction
division and form haploid spores which
functions as the mother cell for gametophytic
phase
 Apospory and Apogamy
• Like pteridophytes,Bryophytes also exhibit
apospory and apogamy
• In apospory, the gametophytes develop from the
vegetative cells of the sporophyte and not from
the spores (Eg. Anthoceros and mosses)
• Aposporous gametophytes resembles normal
gametophyte but possesses diploid number of
chromosomes
• In Apogamy, sporophyte develops from the
gametophyte without gametic fusion (Eg.
Funaria, Phycomitrium etc)
 Classification of Bryophytes
• The term Bryophyta was used for the first time
by Braun (1864) who included algae, fungi,
lichens and mosses in this group
• Eichler (1813) recognized two different classes
Hepaticae
(Liverworts)
Bryophytes
Musei (mosses)
Engler (1892) sub-divided each of these classes in to three
orders in his voluminous work Die Natürlichen
pflanzenfamilien
Order Marchantiales

Class
Hepaticae Order Jungermanniales

Order Anthocerotales

Division Bryophyta
Order Sphagnales

Class Musci
Order Andreales

Order Bryales
• Howe (1899) preferred to elevate order
Anthocerotales to the status of a class and
acocordingly he recognized three classes name
Hepaticae, Anthocerotae and Musci
• Rothmaler (1951) suggested the names
Hepaticopsida, Anthoceropsida and Bryopsida
for Hepaticae, Anthocerotae and Musci
• Prosakeur (1953) supported above classification
and suggested the name Anthocerotopsida for
anthoceropsida, following ICBN
• Prosakeur (1957) proposed a classification
which shows significant deviation from the
above system.
• He included Green algae and all green plants
in the Division Chlorophyta
• Bryophytes are included in this division as a
class, namely Bryopsida
• Bryopsida is further divided in to three sub
classes- Hepaticidae, Anthocerotidae and
Bryidae
 Prosakeur (1957)
Subclass
Class Hepaticidae
Bryopsida
Subclass
Division Anthocerotidae
Chlorophyta
Subclass Bryidae
 Current system of classification
Division Bryophyta
Class
Hepaticopsida Anthocerotopsida Bryopsida

Sub-Class Sphagnobrya Andreaobrya Eubrya

Sphaerocarpales
Marchantiales
Metzgeriales Order
Jungermanniales
Calobryales
Takakiales Anthocerotales Sphagnales Andreales 15 Orders
Funariales
Polytrichales
 “Liverworts”
Includes 6 orders, 9 families, nearly 225 genera
and 8,500 species
• Dorsiventrally flattened gametophyte-Thalloid or foliose
• In foliose forms, leaves are devoid of mid-rib, and are
arranged in 2 or 3 rows on the axis
• Rhizoids- smooth & tuberculate; scales present
• Sex organs develop from superficial cells- mostly dorsal;
very rarely terminal
• Sporophyte may be simple, or differentiated either in to
capsule and foot or in to capsule, foot and seta
• Sporophyte completely depends upon the gametophyte
for its nutritional requirements
• Sprogenous tissue develops from the endothecium of
sporogonium
• The wall of sporogonium is one to several layered and is
devoid of stomata
• Dehiscence of sporogonium is irregular