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Fig.11.19.Frullania Sporophyte L.S.

Spore Dispersal: The wall of the spore capsule is two to several cell layers thick and the walls
of the cells develop bands of thickening. The spores are produced by meiosis and so are haploid.
As the ripe capsule dries out, these bands of thickenings, which are deposited in very specific
patterns, create stresses in the shrinking tissues which cause the capsule to rupture and to split
into four valves (above right). In most leafy liverworts, the spores are discharged by a water
rupture mechanism. Interspersed with the spores are swollen, elongated cells called elaters.
These clear, water-filled elaters have a double spiral band of wall-thickenings on the inside of the
cell wall. The elaters may be free or anchored to the inside walls of the spore capsule
(sporangium) in liverworts, but are anchored at their base in this mechanism.
In some leafy liverworts, such as Frullania, each elater spans the sporangium and is firmly
attached to the sporangium floor at one end and to the roof at the other.
As the four valves of the capsule bend back, as the capsule dries and opens, the elaters are
momentarily stretched and each has a single spiral of thickening and is essentially a stretched
spring in the wall of a water-filled tube. In less than a second the elaters are torn free at their
lower ends, which moves in an arc (as when a stretched spring is bent backwards and then
released from one end) flinging out the spores. In Frullania fragifolia, the large antical lobes of
the bilobed leaves detach, leaving only the smaller postical lobes.

9.6.2-Porella
Division – Bryophyta
Class – Hepaticopsida
Order – Jungermanniales
Family – Porellaceae
Genus – Porella

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External Structure of Porella: The gametophyte of Porella is flat, dorsiventral and foliose.
The prostrate stem or axis is bi- or tripinnately branched. There are three rows of leaves arranged
on the stem, two dorsal rows and one ventral row. The dorsal rows form the lateral leaves and the
leaves of the ventral row are called amphigastria. Leaves are devoid of midribs.
The dorsal leaves are bilobed. The upper anterior lobe, called antical lobe, is larger, usually ovate
with rounded apex; while the lower posterior lobe, called postical lobe or lobule, is much smaller
and narrower with acute apex. The dorsal leaves are closely overlapping and show incubous
arrangement i.e. the lower edge of each leaf is covered by the upper edge of the next leaf below.
This arrangement can be seen when viewed from the dorsal side.
A large number of scattered rhizoids of smooth-walled type arise from the ventral surface of the
stem. The main function of rhizoids is to attach the thallus to the substratum. The absorption of
water and minerals is taking place primarily through the leaves and stem.

Internal Features of Porella:

Stem: The T.S. of the mature stem shows two distinct regions, the cortex and the medulla. The
cortex is 2-3 layered zone, consists of thick-walled parenchymatous cells, while the medulla is
composed of thin-walled, elongated cells.

Leaf: The leaves are very simple in configuration. Each leaf is composed of a single layer of
isodiametric parenchymatous cells containing many chloroplasts. In a few species, leaves contain
oil cells.

Fig.9.20. Porella: (A) Stem T.S. (B) Leaf T.S.

Reproduction:
1. Vegetative Reproduction in Porella: Porella reproduces vegetatively by the following two
methods:
(a) By Progressive Death and Decay of the Gametophyte: The apical growth of the thallus is
accompanied by the progressive death and decay of the older parts of the thallus and consequent
separation of the younger parts at the point of dichotomy of the thallus. The separated branches
develop to form new plants.
(b) By Gemmae: In some species (P. rotundifolia) discoid multicellular gemmae are produced
on the lower surface of the leaves. Gemmae germinate to produce new plants.

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2. Sexual Reproduction in Porella: Porella is dioecious. Male gametophytes are

comparatively smaller than the female gametophyte.
Antheridium: Antheridia are borne on specialized lateral antheridial branches which project
out at right angles to the main axis. The dorsal leaves, called bracts, are smaller than those on the
main branch and are closely imbricated. The ventral leaves (amphigastria) of the antheridial
branch are known as bracteoles. A single antheridium is borne in the axil of each leaf.

Development of Antheridium: An antheridial initial, situated at the base of the young bract,
divides transversely to produce an outer cell and a basal cell. The basal cell does not divide
further and forms the embedded part of the stalk. The outer cell functions as antheridial mother
cell which, by transverse division, forms an upper primary antheridial cell and a lower primary
stalk cell.
A two-celled thick long stalk is developed from the primary stalk cell, following repeated
transverse and vertical divisions. The primary antheridial cell forms the main body of the
antheridium. It forms two identical antheridial cells by a vertical division.
Each of these cells, by a periclinal division, forms two unequal cells, the outer smaller first jacket
initial and the inner larger primary androgonial cell. The latter again divides to form a second
jacket initial. Both the jacket initials form a single layered jacket of the antheridium following
periclinal divisions.

Fig.9.21. Porella: Antheridia Developmental Stages

The primary androgonial cell forms a large number of rectangular androgonial cells following
several divisions in all possible planes. Androgonial cells further transformed to androcyte
mother cell. Each androcyte mother cell following a diagonal division forms two androcytes,
each of which metamorphoses into a biflagellate antherozoid.

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Structure of Mature Antheridium: The mature antheridium is differentiated into a globose

body and a long stalk. The jacket is single-layered at the upper part, while it becomes 2-3 layered
at the lower region through periclinal divisions. The jacket contains inside a mass of androcytes
which ultimately metamorphose into biflagellate antherozoids. The distal part of antheridial
jacket is single- layered and thin. This part of the jacket breaks up into many irregular lobes. This
allows the antherozoids to release into the water.

Archegonium: Archegonia are produced at the apex of archegonial branch on the female plant.
The archegonial branch is much smaller than the vegetative branch, bearing a number of large
perichaetial leaves (bracts). The lower bracts form involucre, while the two upper bracts coalesce
to form a perianth. Ten to fifteen archegonia develop within the perianth.

Development of Archegonium: The archegonia develop in acropetal succession. Each

archegonium develops from a single superficial cell, the archegonial initial which increases in
size and appears as a papillate outgrowth.
The pattern of development of archegonia is identical with that of Riccia and Marchantia.

Structure of Mature Archegonium:

A mature archegonium differentiates into a neck and a venter. The neck is long, comprises of
five vertical rows of neck cells enclosing 6-8 neck canal cells. The venter is 2-layered, consists
of a small ventral canal cell and a large egg. A rosette of 4 cover cells is present at the top of the
archegonial neck.

Fig.9.22.Mature Archegonium of Porella

Fertilization: Like other bryophytes, water is essential for release of sperms and eventual
fertilization in Porella. The process of fertilization is found to be similar with that of other
bryophytes.

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Sporophyte: The zygote increases in size and secretes a wall around itself. Like other
bryophytes; zygote divides transversely to form an epibasal cell and a hypobasal cell. The
hypobasal cell does not divide further, it forms a suspensor. The epibasal cell divides
transversely to form two daughter cells.
These daughter cells undergo repeated transverse as well as vertical divisions in regular sequence
to form an irregular mass of cells. Later, the peripheral amphithecium and inner endothecium are
differentiated by the periclinal divisions in the upper part of the embryo.
The amphithecium gives rise to the capsule wall, while the entire endothecium functions as
archesporium. The archesporium forms the sporogenous tissue through the repeated divisions in
all possible planes.
The sporogenous tissue differentiates into spore mother cells and elater mother cells. The elaters
mother cells become elongated endowed with two spiral thickening and form the sterile elaters.
The spore mother cells divide meiotically to produce haploid spores. The foot and seta develop
from the lower part of the embryo.

Fig.9.23. Sporophyte of Porella (A) Embryo (B) Young Embryo (C) L.S. Mature Sporophyte

Structure of Mature Sporophyte: The mature sporophyte of Porella differentiates into three
parts, viz., foot, seta and capsule. The young sporophyte is enclosed by three protective
coverings — calyptra, perianth and involucre.
(i) Foot: It is the expanded bulbous mass of cells at the base of the sporogonium.
(ii) Seta: It is an elongated structure which connects -the foot with the capsule, made up of
parenchymatous cells.
(iii) Capsule: It is a globose structure containing numerous spores and elaters. The jacket is 3-4
cells thick, made up of thick-walled parenchymatous cells, except for four vertical rows of thin-
walled cells that demarcate the vertical lines of dehiscence.

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Dehiscence of Capsule: At maturity, the seta elongates suddenly, pushing the capsule out of
the calyptra and perianth. As the capsule wall dries up, the capsule now splits into four valves
along the line of dehiscence. The hygroscopic movement of elaters helps in discharging of
spores.

Young Gametophyte: The spore is the first stage of gametophytic generation. A spore has two
concentric walls: the outer ornamented exine and the inner thin- walled intine. Sometimes a third
layer, called perinium, is found outside the exine. The spore germinates immediately after the
fall on the suitable substratum. It differentiates into an apical cell. Then a multicellular thalloid
structure is formed.

Fig.9.24.Gametophyte: Multicellular Thalloid Structure

The rhizoids develop from the lower side and leaves are produced on the upper side.The
germination of spores may take place while the spores are still within the capsule.

Fig.9.25. Life Cycle of Porella

9.7 MORPHOLOGY, ANATOMY AND REPRODUCTION OF

METZGERIALES

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Metzgeriales is an order of liverworts. The group is sometimes called the simple thalloid
liverworts: "thalloid" because the members lack structures resembling stems or leaves, and
"simple" because their tissues are thin and relatively undifferentiated. All species in the order
have a small gametophyte stage and a smaller, relatively short-lived, spore-bearing stage.
Although these plants are almost entirely restricted to regions with high humidity or readily
available moisture, the group as a whole is widely distributed, and occurs on every continent
except Antarctica.

Members of the Metzgeriales typically are small and thin enough to be translucent, with most of
the tissues only a single cell layer in thickness. Because these plants are thin and relatively
undifferentiated, with little evidence of distinct tissues, the Metzgeriales are sometimes called the
"simple thalloid liverworts".
There is considerable diversity in vegetative structure of the Metzgeriales. As a rule, simple
thalloid liverworts do not have structures resembling leaves. However, a few genera, such
as Fossombronia, and Symphyogyna, are "semileafy" and have a thallus that is very deeply
lobed, thus giving the appearance of leafiness. The several semileafy groups within the
Metzgeriales are not closely related to each other, and the currently accepted view is that the
leafy condition evolved separately and independently in each of the groups where it occurs.
Members of the Metzgeriales also differ from the related Jungermanniales in the location of
their archegonia (female reproductive structures). Whereas archegonia in the Jungermanniales
develop directly from the apical cell at the tip of a fertile branch, archegonia in the Metzgeriales
develop from a cell that is behind the apical cell.As a result, the female reproductive organs, and
the sporophytes that develop within them, are always located on the dorsal surface of the
plant. Because these structures do not develop at the apex of the branch, their development in the
Metzgeriales is described as anacrogynous.
The characteristic features of this order are as follows:
1. The gametophyte may be thalloid or differentiated into stem and lateral leaves.
2. In most cases the gametophytes are without internal differentiation of tissues but certain
genera have a central strand of thick-walled cells.
3. The ventral surface of a gametophyte bears smooth-walled rhizoids.
4. The sex-organs are found to be scattered on dorsal surface of thallus.
5. The archegonia arise from the young segments cut off by the apical cell.
6. The mature sporophytes lie some distance back from the growing apex of a gametophyte.
7. The sex organs (antheridia and archegonia) are produced on any branch of the gametophyte or
only on special branches.

9.7.1-Pellia
Division : Bryophyta
Class : Hepaticopsida

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