Life History of Polytrichum

Dr. Habibur Rahman

Associate Professor
J. N. College, Boko
Occurrence
Polytrichum have worldwide distribution with 111
species. In India only 5 species are found and common
species is P. alpinum. They are very common in cool
temperature and tropical regions. Plants live in cool and
shady places. They may be found growing on open soil
or tree trunk as epiphytes.
Gametophytes
The main plant body is gametophyte. The adult plant consists of two parts:
rhizome and upright leafy shoot.
1. Rhizome: It is horizontal portion and grows underground. It bears three
rows of small brown or colourless leaves. It also bears rhizoids. The cells are
rich in protoplasm and oil globules.
2. Upright leafy shoot: The leafy shoots are much longer. It is the most
conspicuous part of the plant. It arises from rhizome. These branches consist of
central axis. These branches bear large leaves arranged spirally.
3. Leaves: Leaves have broad bases. Leaves in the upper portion are green. But
the lower ones are brown. Each leaf has a broad. colourless sheathing leaf base
and narrow distal limb. The mid-rib forms the major part of the leaf. These
leaves possess extra photosynthetic tissue in the form of closely set vertical
plates of green cells. These are known as lamellae. Green lamellae act as
additional photosynthetic tissue.
 Internal Structure
Polytrichum have complex internal structure.
Leaf:
The mid-rib region is thick. But the margins are only one cell thick. The lower
surface is bounded by epidermis. One or two layers of sclerenchymatous
tissues are present above the epidermis. The central tissue of leaf is composed
of thin-walled parenchymatous tissues. Above this are again sclerenchymatous
cells. The upper surface is formed of a layer of large cells from which arise
numerous lamellae. This upper portion is the main photosynthetic region of the
leaf.
Stem: The T.S. of stem shows three regions: medulla, cortex and epidermis.
The medulla is again differentiated into two zones: central zone and peripheral
zone. The cortex consists of thick-walled cells. The innermost layer of cortex
around the conducting strands is known as a mantle. Its cells contain starch
grain. Epidermis is present over the cortex.
Reproduction
Vegetative reproduction:

Vegetative reproduction takes place by following methods:

I. Protonema: The spores germinate to form protonema. Several buds
grow on the protonema. Each bud by of its apical cell develops into
gametophyte.
2. There are also vegetative buds which are found on the rhizoids.
3. Fragmentation: The rhizome gives rise to erect leaf & shoots at
intervals. Death or breaking of shoots separates the erect branches.
These branches behave as independent plants.
Sexual reproduction
Polytrichum is dioecious. Antheridia archegonia occur on different plants.
Antheridial head:
The antheridia are borne in the axillary clusters at the tips of leafy
stems. They are surrounded by a rosette of leaves called perigonial
leaves. These leaves are different from the ordinary vegetative leaves. The
perigonial leaves are spirally arranged. The antheridia are produced in
groups in the axils of these leaves. Thus the antheridial head have different
antheridial groups. Paraphyses also occur among the antheridia.
Mature antheridium is club-shaped. It is composed of a short stalk
and a club-shaped body. Jacket is present around the capsule. Inside the
jacket are present androcyte mother cells. They give rise to biflagellate
sperms.
Development of antheridium
1. The antheridia arise from the embryonic cells at the tip of male
shoot. The embryonic superficial cell forming antheridium is called antheridial initial. It
increases in size. It undergoes transverse division to form lower primary stalk cell and the
upper antheridial mother cell.
2. The primary stalk cell forms a few stalk cells. The antheridial mother cell divides to form
an apical cell with two cutting faces. The apical cell cut off 3-4 segments. Now this apical
cell functions as the operculum cell.
3. The last segment divides by two vertical divisions. It forms peripheral jacket initials and
central primary androgonial cells.
4. Die jacket initials further divide to form a single-layered jacket. The primary androgonial
cells divide to form androgonial cells.
5. The last generation of primary androgonial cells is called the androcyte mother cells. Each
androcyte mother cell gives rise to two coiled biflagellate sperms.
6. The antheridia always dehisce in the presence or wren The operculum cell is thrown out
and pore is formed at the apex. Sperm mass contained in mucilage comes out.
Archegonial head:

The flask-shaped archegonia are borne at the apices of leafy

stems. Archegonitim is surrounded by perichaesial leaves. These
leaves overlap to form a closed bud-like structure. The archegonia
occur in cluster of 3 to 6.
Mature archegonium is flask-shaped. It has a thick
multicellular stalk. The neck is long and twisted. It contains neck
canal cells. The neck consists of 6-vertical rows of cells. Neck
gradually merges into venter. Venter contains upper small venter
canal cell and lower large egg cell. Paraphyses are absent.
Development of archegonium
 Development of archegonium
1. Any apical cell in the apical region acts an archegonial initial.
The archegonial initial enlarge. It divides by a transverse division to form
lower primary stalk cells and upper archegonial mother cell.
2. The primary stalk cell forms a massive stalk. The archegonial mother
cell forms the main body of archegonium. It undergoes three vertical
division s to form three peripheral cells surrounding an axial cell.
3. Three peripheral cells divide to form 2-3-layered jacket around the
venter. The axial cell divides transversely to form inner central cell and
outer apical cell.
4. Central cell forms upper small venter canal cell and lover large egg
cell. Apical cell divides to form long neck which consists of 6 vertical
rows of cells. The cells cut off from the base foem neck canal cells.
Fertilization:

The sex organs dehisce in the presence of water. The venter

canal cell and the neck canal cells dissolve to form mucilage. This
mucilage exerts pressure and the neck opens out. The mucilage comes
out of the neck. The sperms reached the archegonial heads by rain
water. They are attracted towards the archegonia. One of the sperm
swims down the open neck and reaches the base. It fuses with the egg
to form oospore. Oospore is the first stage of sporophytic generation.
Sporophyte
Development of Sporogonium:
1. The oospore divides transversely to form upper (epibazal) and lower (hypobasal) cell.
2. The hypobasal region forms foot and lower part of seta. The foot region consists of thin-
walled cells. It is embedded ir the stalk of the archegonium. The cells of the seta are larger
and poor in cytoplasmic contents.
3. The epibasal region forms upper portion of seta and the capsule. Epibasal cell divides to
form young embryo. Young embryo is cylindrical and completely surrounded by calyptra.
Cells of the embryo divide to form amphithecium and the endothecium regions. 8-
amphithecium cells are surrounded by a group of 4- endothecium cells.
4. Endothecium forms central conducting strands of apophysis. It forms columella and
spore sac of theca. It also forms membranous tissues of the operculum. The outermost layer
of endothecium forms archesporium or spore mother cells. These cells divide meiotically to
form haploid spores.
5. The amphithecium divides to form seven rings of cells. These cells give rise to spongy
tissues and epidermis of apophvsis. They also form outer wall of theca.
Structure of Mature Sporogonium:
(Sporophyte)
The mature sporogonium is differentiated into
foot, seta and capsule.
Foot.: The foot is buried deep in the tissue of
gametophyte. It is absorptive in function. It
consists of thin-walled narrow cells containing
dense cytoplasm.
Seta: The seta is several inches long. It carries
the capsule high into the air. It also conducts
water and food. It consists of epidermis, cortex
and central conducting strands.
Capsule: The upper part is capsule. It is
differentiated into three regions: apophysis,
theca and operculum.
Dispersal of spores:
Cells of the epiphragm dry up during dry conditions. It separates the
operculum. The calyptra falls. Epiphragm also dries up between the peristome
pores. The central tissue of theca region except the spores degenerates. Thus
the spores lie free in the centre of the capsule at maturity. Spores come out
through pores. They are dispersed by wind.
Structure and germination of spores:
The spores are yellow. Each spore is uninucleate and has two wall laye:s. The
outer layer is exosporium (exine). The inner layer is endosporium. The spore
germinates under favourable conditions.
Exosporium ruptures and endosporium comes out. It forms prntonema.
Protonema develops many buds. These buds produce new moss plants.
 Altrnation of generation

Polytrichum shows heteromorphic alternation of generation.

Gametophyte: The plant body is gametophytes. Gametophyte is haploid.
It develops antheridia and archegonia. Antheridia produce antherozoids
and archegonium produces egg. Antherozoids fuse with egg to produce
diploid oospore.
Sporophyte: The oospore is the first stage of sporophyte generation. It is
diploid generation. Sporophyte has three parts: foot, seta and capsule.
Haploid spores are produced in the capsule by meiosis. Spore is the first
stage of gametophyte. Spores germinate to produce protonema stage. It
gives rise to mature gametophyte completing the life cycle.