18BZO14A-U2

CLASS: OOMYCETES
ORDER: PERSANOSPORALES
FAMILY: ALBIGINACEAE
GENUS: ALBUGO CANDIDA

Mycelium of Albugo:
It is well developed and consists of branched, aseptate, coenocytic hyphae. The hyphae live
and ramify in the intercellular spaces of the susceptible host tissue. The hyphal wall
contains cellulose and not chitin. The hyphal protoplasm is granular and vacuolate in the
older parts.

It contains numerous nuclei, oil globules and glycogen. Electron micrographs reveal the
presence of mitochondria, endoplasmic reticulum and ribosomes.

The cytoplasmic membrane which is closely appressed to the hyphal wall forms
lomasomes. Septa remain suppressed in the actively growing hyphae but appear to separate
reproductive structures and to seal off injured parts.

The fungus mycelium grows vigorously. The hyphae branch and ramify within the host
attacking the tissues adjoining the point of infection. Sometimes both Albugo and
Peronosporaoccur on the same host particularly Capsella bursapastoris. Albugo can,
however, be distinguished from Peronospora by the smaller diameter of its hyphae and
more numerous, vesicular haustoria (Fig. 6.50).
The intercellular hyphae of this obligate parasite produce intracellular haustoria in the
mesophyll cells of the host (Fig. 6.50). The haustorium arises as a lateral outgrowth at the
site where the hyphal wall is tightly pressed against the mesophyll cell wall.
An electron- dense amorphous material known as the penetration matrix is usually
deposited at the site of contact between the host and the hypha cell walls. It is described as
the penetration site (Fig.
6.51).

The slightly crescent-shaped bulge of the haustorial mother cell known as haustorial initial
perforates the host cell wall at the penetration site and protrudes into the lumen of the
mesophyll cell to develop into a haustorium. It is bordered by the invaginated host plasma
membrane.

With light microscope the haustorium is seen as a small, spherical structure consisting
of two parts namely:
(i) The haustorial stalk or neck and

(ii) The terminal haustorial head or body.

The haustorial stalk passes through the penetration site to connect the haustorial body to
the hyphal wall in the intercellular space between the mesophyll cell. Usually one or two,
sometimes more, haustona are seen in the thin peripheral layer of the host cell cytoplasm
adjacent to the chloroplasts.

Reproduction in Albugo:
Asexual Reproduction in Albugo:
When the mycelium has reached a certain stage of maturity it epidermis produces pads of
hyphae at certain areas just below the epidermis. The tips of hyphae constituting the mat
grow verticlly into short, upright, thick-walled, unbranched club-shaped hyphae.

These are the sporangiophores (=conidiophores). They are arranged in a closely packed
palisade like layer forming a sorus between the epidermis and the mesophyll of the host
leaf. Each sporangiophore appears constricted at its junction with the subtending hypha.

The lower two-third portion of sporangiophore is narrow, thick-walled, with a undulating

surface whereas the upper one-third is broader, thin-walled with a smoother surface.
According to Khan (1977), the sporangiophore wall towards its proximal end consists of
two layers, the outer more electron-dense and thicker than the inner layer.

(a) Abstriction of sporangia:

In the lower fungi (Phycomycetes) Albugo is unique in that its lemon- shaped sporangia
are produced in basipetal chains at the tips of clavate sporangiophores. Two different views
have been put forth to explain their mode of development.
According to one view, the sporangial chains in Albugo are abstricted by percurrent
proliferation. The second view advocates the blastic mode of development.

(i) Percurrent Proliferation (Fig. 6.53):

The sporangia in Albugo which are cut off in succession are arranged in a basipetal chain
on the sporangiophore. According to Hughes (1971), they are produced by successive
proliferations of the sporangiophore subtending a sporangium.

This mode of development of sporangia is termed per-current proliferation. Generally the

sporangiophore increases in length as each successive sporangium is cut off from each
successive proliferation at a higher level than the previous one. The first formed
sporangium is a aleuriosporangium.

Reaching a certain size it is delimited from the sporangiophore by a basal septum. The
latter eventually splits into two halves so that the subsequent proliferation of
sporangiophore involves the exposed half septum. In Albugo each successive sporangium
is capable of seceding from the sporangiophore or from the young sporangium.

The second sporangium is thus formed by proliferation of the sporangiophore with total
involvement of the half of the fractured transverse septum exposed by the seceding first
sporangium above it. Apart from this, septum is seen at the apex of the young sporangium.

The second sporangium is delimited in the same manner as the first. As the second
sporangium increases in size it pushes the first upward without disjunction. The process is
repeated resulting in a chain of sporangia. Probably the septum seen at the apex of each
younger sporangium thickens on both sides to form a connective between the successive
sporangia in the chain.
According to Hughes, besides increase in length of sporangiophores, this method of
sporangium development is accompanied by marked lamination and thickening of the
walls of the sporangiophores. The mature sporangiophores are thus longer, more thick-
walled and show annellations.

Thakur (1977) corroborated findings of Hughes (1971) on formation of sporangia by

percurrent proliferation in Albugo.

(ii) Blastic mode of Sporangium development (Fig. 6.54):

According to Khan (1977) the sporangiophore has a fixed sporogenous locus at its apex.
The sporangial initial arises as a bud from it (A). It contains about 4-6 nuclei and dense
cytoplasm.

The two wall layers of the sporangium initial are continuous with those of the
sporangiophore wall. Reaching a certain size, the initial is delimited by a basal septum near
the sporogenous locus. It becomes the first sporangium and the oldest in the chain (B).

The septum is formed by the centripetal growth of the inner layer of the sporangiophore
wall (C). A nearly complete septum has a narrow central canal and consists of three layers,
upper and lower electron dense and the thick middle one of less electron density (D).

After the completion of the basal septum and conversion of the initial into a full-fledged
sporangium, a new sporangium initial grows as a bud from the sporogenous locus (B). It
pushes the newly-formed sporangium upward. Thus only one sporangium is formed at a
time.

As the second sporangium initial grows to the normal size, it is also delimited by the
formation of a basal septum as the first. The repetition of the process results in the
formation of a basipetal chain of sporangia. Soon after the formation of the first
sporangium, the breakdown of its basal septum begains.

It is the middle layer which starts disintegrating from its periphery inwards whereas its
upper layer fonns the wall of the upper sporangium and the lower layer completes that of
the lower sporangium (E-F).

The fibrous product of dissolution of the middle layer is held in position by the pellicle
which covers both the sporangia and the sporangiophore. It is seen as a connective or
disjunctor between the successive sporangia in the chain. Khan (1977) did not notice any
increase in length of the sporangiophore during sporangia formation nor did he observe any
annellations on the sporangiophore surface.

Sporangia (Fig. 6.55 A, B):

They are small, hyaline, nearly spherical or lemon- shaped structures with a smooth or
somewhat punctate surface.

The basipetal arrangement of sporangia in the chain (with the oldest at the top and the
youngest at the base of the chain) serves two useful purposes:
(i) It permits ready dispersal of the oldest sporangia by air currents or rain water, and

(ii) It helps in the proper nourishment of the younger ones.

(i) Ultrastructure of sporangium (B):

According to Khan (1977), each newly formed sporangium is lemon-shaped and is about
19- 22 by 14-17 µ m in size. It bears remnants of the connectives or disjunctor pads at both
the ends. The sporangium wall is differentiated into two distinct layers. The outer is more
electron dense than the inner.

Within the sporangium wall is the highly convoluted plasma membrane enclosing the
dense cytoplasm containing up to 4 nuclei. Besides, the cytoplasm contains endoplasmic
reticulum, mitochondria, perinuclear, dictyosomes, ribosomes both free and attached to
endoplasmic reticulum, vesicles of various kinds and lipid droplets.

Towards maturity the sporangial wall especially, its inner layer increases in thickness and
the number of lipid droplets decrease as the sporangia matured. The oldest sporangia have
none.

The endoplasmic reticulum becomes accumulated in the peripheral cytoplasm. Towards the
end of sporangial maturation, the dictyosomes become quiescent, mitochondria decreased
in number and also the amount of endoplasmic reticulum. The sporangial wall increased 3-
fold the thickness.

(ii) Dispersal of sporangia:

The chains of sporangia lengthen and press on the epidermis above. This causes the leaf
surface to bulge. The overlying epidermis eventually bursts over the growing sporangial
sorus and exposes the white shining pustules consisting of masses of sporangia.

The pustules look like white blisters. The exposed sporangia are white. The distal ones by
this time have matured. As the sporangia mature the connectives or gelatinous pads
between them dry, shrink and finally disintegrate in moist air. The sporangia in the chain
thus separate. They are then blown away in the air by wind or washed away by rain water.
Germination of sporangia (Fig. 6.56):
Landing on a suitable host the sporangia begin to germinate within two or three hours
under suitable conditions.

At the time of germination they behave in either of the following two ways depending on
temperature conditions:—
Indirect Germination (Fig. 6.56 B-E):
In the presence of moisture and low temperature, the sporangium functions as a
zoosporangium (B). The optimum temperature for germination of sporangia is 10°C. It
absorbs water and swells. A few vacuoles appear in its granular cytoplasm.

Later the vacuoles disappear and the multinucleate protoplast undergo division. It divides
to form five or eight polyhedral uninucleate daugher protoplasts. Meanwhile an obtuse
papilla forms on one side of the sporangium.

Each daughter protoplast shapes into a slightly concave-convex zoospore (E). It has a disc-
like contractile vacuole on one side and is furnished with two flagella, one short and one
long. The former is of tinsel type and the later whiplash. The flagella are attached laterally
near the vacuole.

As the zoospores are differentiated, the papilla swells and opens. The zoospores still
immobile, emerge usually one by one (Fig. 6.56 C). According to Vanterpool, the
zoospores are, at first, released in a sessile vesicle formed by the swelling of the papilla.
The vesicle soon vanishes.
Germination of zoospore and infection of host (Fig. 6.56 F-H):
Moisture on the surface of the host is essential for germination and infection. The released
zoopores swim about in water for a while (E). Finally they settle down on the host, retract
the flagella and round off.

Each secretes a wall around it (F). The encysted zoospore (cyst) then germinates. It puts
out a germ tube (G) which gains entrance into the host thought a stoma (H). Once within
the host tissue the germ tube grows and forms the mycelium

(ii) Direct Germination (Fig. 6.56 I-J):

At high temperature and under comparatively dry conditions the sporangium behaves like a
conidium (I). It germinates directly to form a germ tube (J). The conidial method of
germination of sporangia in Albugo is, however, not common.

The germ tube penetrates the host through a stoma or, through an injury in the epidermis.
Within the host it develops into a mycelium. Re-infection of the host and infection of other
healthy plants in the vicinity goes on by the production of sporangia throughout the
growing season.

Sexual Reproduction in Albugo:

It is oogamous:
The male sex organ is called an antheridium and the female oogonium (A). They are
developed near each other in the intercellular spaces of host tissues towards the end of the
growing season. When the mycelium ages, some hyphae grow deep and lie buried in the
intercellular spaces of the tissues of the stem, or petioles.

The sex organs arise on separate hyphae called the male and the female hyphae (A). The
two soon establish contact. The antheridium comes in contact with the oogonium at the
side. The development of sex organs within the host tissue is externally indicated by
hypertrophy and distoration in shape in the particular organ.

(a) Oogonium:
It arises as a globular enlargement of the tip of the female hypha. Sometimes the oogonium
is intercalary in position. The swelling is multinucleate(A) Across wall appears below this
inflation (B). It separates the terminal oogonium from the rest of the female hypha. The
young ooglum has highly vacuolated contents.

The numerous nuclei and vacuoles are evenly distributed and the usual cell organelles are
dispersed throughout the oogonium. The nuclei divided mitotically and increase in number
as the oogonium advanced towards maturity. After the first Sc division the ooginial
cystoplasm shows marked zonation (C).
It becomes differentia into two distinct regions with the rearrangement of the numerous
nuclei and other cellular organellers. Most of the original cytoplasm of the oogonium forms
the central,rounded dense ooplasm. It is multinucleate and contains only a few
mitochondria,ribosomes and cisternae of ER. It is rich in lipid vesicles and reserve vesicles
containing electron dense inclusions (reserve globules).

The ooplasm is surrounded by the peripheral cytoplasm constituting the periplasm. It is

more vacuolate and spongy. The vacuoles are large. Besides, the periplasm is rich in
nuclei, mitochondria endoplasmic reticulum and ribosomes. It has protoplasm of thinner
consistency. Sometime after all the nuclei of ooplasm migrate into the periplasm (D) and
become arranged in a ring.

Here they divide mitotically with the spindles lying in such a way that one pole of each that
one pole mitosis one daughter nucleusthe ooplasm and the other in the periplasm (E). At
the end of mitosis one daughter nucleus of each spindle goes to the ooplasm and the other
to the periplasm (F). However, the ooplasm at maturity has a single centrally located
nucleus (G).

There are two views with regard to this uninucleate condition of the mature ooplasm.
According to one view, all the nuclei excepting one are extrude from the ooplasm and are
deposited in the periplasm. The second view IS that in the later stages of development all
the nuclei in the ooplasm excepting one degenerate and disappear. The uninucleate
ooplasm functions as the female gamete or egg or oosphere (G).

(b) Antheridium:
It is an elongated club-shaped cell (A). It is multinucleate. The antheridium is developed at
the end of a male hypha lying close to the oogonium. The end of the male hypha enlarges
into club-shaped swelling.

The latter is then cut off by a cross wall from the rest of the male hypha (B). This terminal
dub-shaped cell is called an antheridium. It contains several nuclei (usually 6-12), but only
one is functional. The paragynous antheridium comes in direct contact with the oogonium
at the side (C).
(c) Fertilisation:
At the point of contact of antheridium with the oogonium, the walls become very thin. A
portion of the contents of the oogonium surrounded by a thin membrane grows into a
papilla-like outgrowth (G).

This papilla-like oogonial bulging is called the receptive papilla It is functionless. The
receptive papilla bulges (G) into the antheridium but soon disappears. This is followed by
the formation of a slender tubular outgrowth from the antheridium.

It is the fertilisation tube (H). The fertilisation tube passes through the thin spot in the
oogonial wall and enters the multinucleate periplasm. It then dips deep into the ooplasm.
Prior to this a spherical and granular cytoplasmic body appears in the centre of the
oosphere (H). It is known as the coenocentrum.
The single functional female nucleus is attracted towards it and becomes attached to a point
near it.

The fertilisation tube finally reaches the coenocentrum and ruptures (I) at the tip to
introduce a single male nucleus which fuses with the female nucleus. Thereafter the
fertilisation tube collapes but persists and the coenocentrum vanishes. On removing or
displacing the attached antheridium Tewan and Skoropad (1977) observed a clear hole
surrounded by some fibrous material.
Oospore (Fig. 6.58 A-C):
Tewari and Skoropad (1977) investigated the fine structure and development of A. Candida
oospores. According to them, the young oospore is delimited from the vacuolate periplasm
by an electron-dense cell wall.

The single layered cell wall of the young oospore encloses dense cytoplasm containing a
group of reserve vesicles, lipid vesicles and a few membranous organelles. It is surrounded
by periplasm rich in vacuolate cytoplasm containing membranous organelles.

Further development of oospore if marked by the deposition of 4 layers, two on the outer
and two on the inner side of the first (original) layer of the young oospore. The mature
oospore thus has a thick highly differentiated 5-layered wall.

External to the oospore wall are the two addition protective investments formed by the
persistent periplasm and the oogonial wall. The thick highly differentiated oospore wall
together with the two surrounding additional layers contributes to the longevity of Albugo
oospore. The authors suggest that periplasm plays an active part in deposition of oospore
wall layers.
Within the fully developed oospore wall is the scanty cytoplasm surrounding a large central
reserve globule. Some small bodies resembling the reserve globules in appearance and
numerous lipid vesicles occupy most of the space between the oospore wall and the central
reserve globule.

The highly differentiated thick, oospore wall together with the two additional layers
constituted by the persistent perisperm and the oogonial wall provides protection and the
numerous lipid vesicles in the oospore cytoplasm furnish energy for the long dormancy or
overwintering by oospores in Albugo.

The outer layer of the oospore wall is comparatively thicker. It is warty or tuberculate. In
other species it may have a network of ridges or other patterns.

Location of meiosis:
The place of meiosis in Albugo is still under dispute. Stevens (1899) suggested that Albugo
possesses diploid somatic nuclei which undergo meiosis in the gametangia (antheridia and
oogonia). His interpretation was disputed by later workers. They held that Albugo and
other Oomycetes, as a whole, are haploid. Oospore is the only diploid structure in the life
cycle.

It undergoes zygotic meiosis. Walker (1969) observed that after fertilization when a thick
wall is being developed around the oospore, its diploid nucleus divides repeatedly to form
32 nuclei. Probably the two earlier of these divisions constitute meiosis. In this 32 nucleate
stage, the oospore enter the resting stage and tides over the period unfavourable for growth.
However, Sansome and Sansome (1974) have advanced evidence in support of
gametangial meiosis and diploid life cycle in Albugo Candida. They hold that the first two
divisions of nuclei which occur in the gametangia constitute meiosis. This view is gaining
ground.

Germination of Oospore (Fig. 6.59):

On the onset of conditions favourable for growth, the oospore germinates. The central
globule and the lipid droplets gradually disappear. The contents of the oospore assume
uniform granular appearance.

The diploid nucleus undergoes repeated divisions to form many nuclei (about 100 or even
more). A small amount of cytoplasm gathers around each daughter nucleus. Numerous
uninucleate daughter protoplasts thus result. Each of these metamorhoses into a biflagellate
zoosore.

The oospore then germinates to release the zoospores by either of the two following
methods:— (i) The thick oospore wall cracks. A germ tube emerges through the split. It
ends in a thin vesicle. From the oospore the zoospores pass into the vesicle. Soon the
vesicle perishes to liberate the zoospores.

(ii) Through a crack in the oospore wall emerges a thin sessile’ vesicle containing the
zoospores (A).
The vesicle soon bursts to liberate the zoospores.

(iii) Verma and Petrie (1975) described direct germination of oospore without the
intervention of zoospores. In this method after the disappearance of the central globule
and the lipid dropletes the contents of oospore assume uniform granular apearance.

The thick oospore wall then cracks. The contents then emerge in the form of one or two
simple or branched germ tubes. Direct penetration of the host by a germ tube has not been
reported. Good germination occured at 10-20°C. They observed that 71 percent of the
oospores collected from the field in August germinated within two weeks.
Structure of zoospore:
The liberated zoospore is a biflagellate, uninucleate structure which is reniform in shape.
The two unequal flagella arise from a depression on the concave side. The shorter
flagellum is of tinsel type and the longer one of whiplash type. The former is directed
forward and the latter trails behind when the zoospore is in motion.

Germination of Zoospore (Fig. 6.59 D-E):

On coming in contact with a suitable host. The flagella are withdrawn. It then rounds off
and secretes a wall around it (D). Soon the encysted zoospore (cyst) puts out a germ tube
(E) which enters the host tissue through a stoma. Once within the host tissue the germ tube
grows vigorously and forms a new mycelium.

Primary infection of host:

According to Verna et al. (1975), the zoospores produced in the germinating oospores
serve as a primary inoculum for infection of the susceptible host. The emerging cotyledons
are the infection sites.

After penetration the first haustorium originates near the tip of the young hypha. The latter
then continues to grow leaving the haustorium as a side branch. The formation of the first
functional haustorium is the critical step in primary infection. It indicates the establishment
of a compatible functional host-parasite relationship.

From then onwards hyphal growth increases rapidly. The hyphae grow around the palisade
mesophyll cells with haustoria penetrating the adjacent cells. The number of haustoria per
cell varies from one to several. In the course of time a mycelial base is established inside
the host tissue (cotyledons).

It produces masses of zoosporangia on the cotyledons which serve as secondary inoculum

in initiating systemic infection. The parasite ultimately reaches the inflorescence region
where it produces the oospores.
1. Cell Structure of Saccharomyces 2. Vegetative Body of Saccharomyces
3. Reproduction 4. Life Cycle Patterns.
Cell Structure of Saccharomyces:
The genus Saccharomyces (Gr. Saccharon, sugar; mykes, fungus) consists of about
41 species. 5. cerevisiae, commonly known as Brewer yeast or Backer’s yeast is
used widely in wine and baking industry.

It produces two types of enzymes: an extracellular invertase and an intracellular

zymase. The invertase hydrolyses canesugar to dextrose or invert sugar and zymase
breaks invert sugar into ethyl alcohol and carbon dioxide.
Vegetative Body of Saccharomyces:
The thalloid plant body is unicellular, but during rapid multiplication by budding
the cells may remain attached in chain forming pseudo- mycelium (Fig. 4.38). The
cells may be globose, elliptical, oval to even rectangular in shape and measure
about 5-6 x 6-8 µm.

Electron microscopic studies (Fig. 4.34, 4.35) and chemical analysis of

Saccharomyces cerevisiae show that the cells are surrounded by a distinct cell wall
with three layers. The outermost layer mainly consists of protein-mannan and some
chitin; the middle layer mainly of glucan and the innermost layer consists of
proteinglucan.

Some phosphate and lipids are also present, while cellulose is absent in the cell
wall. Inner to the cell watl is the cell membrane (plasmalemma), i.e., an usual unit
membrane having series of shallow, elongated pits or invaginations (Fig. 4.35).

In the centre, the cell having a large central vacuole, limited by a single membrane,
the tonoplast, which contains a watery substance, granules of polymetaphosphate
and lipid.
The cytoplasm is granular and contains organelles like nucleus, mitochondria, golgi
apparatus, endoplasmic reticulum, ribosome and other substances like glycogen
bodies, volutin granules, oil globules etc.

Some hydro- lytic enzymes like proteases, esterases, ribonuclease etc., are also
present in the cytoplasm. The nucleus having outer perforated double unit
membrane remains by the side of the vacuole. The nucleus is bipartite in nature
having major Feulgen positive and a smaller Feulgen negative regions (Moor and
Muhlethaler, 1963).

Reproduction in Saccharomyces:
Saccharomyces reproduces by vegetative, asexual and sexual means.

A. Vegetative Reproduction:
Vegetative reproduction takes place by fission and budding.

(a) Fission:
It takes place during favourable condition. In this process, single vegetative cell
forms two daughter cells of equal size (Fig. 4.36). During fission, a constriction
appears in the middle of the cell and simultaneously nucleus undergoes mitotic
division.

Both the steps progress simultaneously. After nuclear migration, one at each side,
partition wall forms almost in the half way of the mother cell and, as such, two
daughter cells are formed.

(b) Budding:
Budding also takes place during favourable condition. The protoplasm of vegetative
cell swells up at one side in the form of a bud (Fig. 4.37). The nucleus undergoes
mitotic division. Out of two nuclei formed by mitosis, one goes to the bud and
other one remains in the mother. Bud enlarges and eventually cuts off from the
mother by partition wall.

The size of the bud is always smaller than the mother cell. After maturation, these
bud separate from the mother and leave a convex scar on the surface, called bud
scar. Similar scar with concave surface remains on the wall of the bud, called birth
scar.

Sometimes due to rapid division, large number of buds develop without being
detached from one another and persist in the form of branched or unbranched chain,
called pseudo- mycelium (Fig. 4.38). Finally the cells get detached and grow
individually.
B. Asexual Reproduction:
It takes place during unfavourable condition by the formation of thick walled spore,
called endospore (Fig. 4.39). During this process nucleus divides mitotically and
forms four nuclei.
The protoplast divides into four units, each with one nucleus and forms four
endospores.
During favourable condition, endospore germinates by budding and buds grow
individually.

C. Sexual Reproduction:
Sexual reproduction takes place during unfavourable condition. In this process, two
vegetative cells or ascospores behave as gametangia (Fig. 4.40). Two such cells
come very close and develop beak-like outgrowth towards each other. Both the
outgrowths come in contact and the intervening walls between them dissolve.

The nuclei of both the gametangia come to the fused outgrowth (conjugation tube)
and they fuse therein to form a diploid zygote. The zygote behaves as an ascus. The
diploid nucleus of zygote undergoes meiotic division forming 4 or 8 (with
additional mitosis) ascospores. The ascospores are liberated by breaking the ascus
wall and behave as somatic cell.
Life Cycle Patterns of Saccharomyces:
Three patterns of life cycle are found in yeast:
Haplobiontic, diplobiontic and haplodiplobiontic (Fig. 4.41):
1. Haplobiontic Type:
This type of life cycle is characterised by more elaborate haploid phase than the
diploid phase, found in Schizosaccharomyces octosporus (Fig. 4.41 A). The diploid
phase is restricted only in the zygote. The vegetative cells are haploid and behave
as gametangia.

Two such gametangia fuse together and form a diploid cell. The diploid cell
behaves as an ascus whose nucleus divides first meiotically, then mitotically;
results in the formation of eight ascospores. After maturation, the ascospores
liberate by bursting the ascus wall. The ascospores then behave as vegetative cell
and continue multiplication through budding.
2. Diplobiontic Type:
This type of life cycle is characterised by more elaborate diploid phase than the
haploid phase, found in Saccharomycodes ludwigii (Fig. 4.41 B). The haploid
phase is restricted only in ascospore, with short duration. The ascospores behave as
gametangia and, without liberating from ascus, they unite in pair. The paired
gametangia after fusion produce diploid zygote.

The zygote then germinates by producing germ tube which comes out through the
ascus wall. The germ tube becomes multicellular from which diploid sprouts
develop by budding. After detachment from the mother, the diploid sprouts
function as asci and produce four ascospores by reduction division.

3. Haplo-Diplobiontic Type:
This type of life cycle is represented by haploid and diploid phases, of more or less
equal duration, found in Saccharomyces cerevisiae (Fig. 4.41 C). The haploid cells
of opposite mating type normally multiply by budding. Two such cells of opposite
mating behave as gametangia and undergo fusion. The fused gametangia develop a
diploid zygote.

The diploid zygote like the haploid cells undergoes budding and forms many
diploid cells. With the scarcity of food, the diploid cell behaves as an ascus and by
reduction division it forms four haploid ascospores. After liberating from the
mother wall, the ascospores undergo budding and form many haploid somatic cells.

Economic Importance Of Fungi

Fungi are one of the most important microorganisms carrying out essential
functions which may not be visible but are crucial in accelerating biological
processes. These eukaryotic entities are closely related to the fauna and distantly
related to the flora with respect to their characteristics and classification. A few
features that set fungi apart from other microbes are :
• Fungi are osmotrophic – can absorb food
• Possess a characteristic hypha at their tips which carry out food exploration
• They possess nucleic cells containing chromosomes
• They are heterotrophs, cannot synthesize their own food.
• Reproduce through spore formation(Mushrooms)
Although we usually think of fungi as food perishing agents, they are economically
very beneficial. Fungi are extensively used across industries in various forms and
stages of processes.

Fungi are an important organism in human life. They play an important role in
medicine by yielding antibiotics, in agriculture by maintaining soil fertility, are
consumed as food, and forms the basis of many industries. Let us have a look at
some of the fields where fungi are really important.

Importance in Human Life

Fungi are very important to humans at many levels. They are an important part of
the nutrient cycle in the ecosystem. They also act as pesticides.

Biological Insecticides
Fungi are animal pathogens. Thus they help in controlling the population of pests.
These fungi do not infect plants and animals. They attack specifically to some
insects. The fungus Beauveria bassiana is a pesticide that is being tested to
control the spread of emerald ash borer.

Reusing
These microbes along with bacteria bring about recycling of matter by
decomposing dead matter of plants and excreta of animals in the soil, hence the
reuse enriches the soil to make it fertile. The absence of activities of fungi can have
an adverse effect on this on-going process by continuous assembly and piling of
debris.
Importance in Medicine
• Metabolites of fungi are of great commercial importance.
• Antibiotics are the substances produced by fungi, useful for the treatment of
diseases caused by pathogens. Antibiotics produced by actinomycetes and
moulds inhibits the growth of other microbes.
• Apart from curing diseases, antibiotics are also used fed to animals for
speedy growth and to improve meat quality. Antibiotics are used to preserve
freshly produced meat for longer durations.
• Penicillin is a widely used antibiotic, lethal for the survival of microbes.
The reason it is extensively used is since it has no effect on human cells but
kills gram-positive bacteria.
• Streptomycin, another antibiotic is of great medicinal value. It is more
powerful than Penicillin as it destroys gram-negative entities.
• Yield-soluble antibiotics are used to check the growth of yeasts and bacteria
and in treating plant diseases.
• Administration of Griseofulvin results in the absorption by keratinized
tissues and are used to treat fungal skin diseases(ringworms).
• Ergot is used in the medicine and the vet industry. It is also used to control
bleeding post-child-birth.
• LSD – Lysergic acid, is a derivative of ergot and is used in the field of
psychiatry.
• Consuming fungi called Clavatia prevents cancer of the stomach.

Importance in Agriculture
The fungi plant dynamic is essential in productivity of crops. Fungal activity in
farmlands contributes to the growth of plants by about 70%.
Fungi are important in the process of humus formation as it brings about the
degeneration of the plant and animal matter.
They are successively used in biological control of pests. Plant pests are used as
insecticides to control activities of insects. For example – Empausa sepulchralis,
Cordyceps melonhae. Use of fungal pesticides can reduce environmental hazards
by a great extent.
Fungi are also used in agricultural research. Some species of fungi are used in the
detection of certain elements such as Copper and Arsenic in soil and in the
production of enzymes. For instance, biological and genetic research on fungi
named Neurospora led to the One Gene One Enzyme hypothesis.
The fungi live in a symbiotic relationship with the plant roots known as mycorrhiza.
These are essential to enhance the productivity of farmland. 80-90% of trees could
not survive without the fungal partner in the root system.
Importance in Food industry
Some fungi are used in food processing while some are directly consumed. For
example – Mushrooms, which are rich in proteins and minerals and low in fat.
Fungi constitute the basis in the baking and brewing industry. They bring about
fermentation of sugar by an enzyme called zymase producing alcohol which is
used to make wine.
Carbon dioxide- a byproduct in the process, is used as dry ice and also in the baking
industry to make the dough (rising and lightening of dough).
Saccharomyces cerevisiae is an important ingredient in bread, a staple food of
humans for several years. It is also known as the baker’s yeast.

Life Cycle of Marchantia (With Diagram) | Hepaticopsida

The gametophytic phase, reproduction and sporophytic phase in the life cycle
of marchantia.
Gametophytic Phase of Marchantia:
External Features of Gametophyte:
The plant body is gametophytic, thalloid, flat, prostrate, plagiotropic, 2-10 cm. long
and dichotomously branched (Fig. 1 A).

Dorsal surface:
Dorsal surface is dark green. It has a conspicuous midrib and a number of
polygonal areas called areolae. The midrib is marked on the dorsal surface by a
shallow groove and on the ventral surface by a low ridge. Each polygonal area re-
presents the underlying air chamber.

The boundaries of these areas represent the walls that separate each air chamber
from the next. Each air chamber has a central pore. The midrib ends in a depression
at the apical region forming an apical notch in which growing point is situated (Fig.
28 B).

Dorsal surface also bears the vegetative and sexual reproductive structures. The
vegetative reproductive structures are gemma cup and develop along the midrib.
These are crescent shaped with spiny or fimbriate margins and are about one eighth
of a inch in diameter (Fig. I A, 15).

Sexual reproductive structures are borne on special Stalked structures called

gametophores or gametangiophores. The gametophores bearing archegonia are
called archegoniophores and that bearing antheridia are called antheridiophores
(Fig. 1 A, B).

Ventral surface:
The ventral surface of the thallus bears scales and rhizoids along the midrib.
Scales are violet coloured, multicellular, one cell thick and arranged in 2-4
rows (Fig. 1 C). Scales are of two types:
(i) Simple or

ligulate (ii)

Appendiculate

Appendiculate (Fig- 1 C, D) scales form the inner row of the scales close with
midrib. Ligulate scales form the outer or marginal row and are smaller than the
appendiculate scales (Fig. 1 C, E).

Rhizoids are unicellular, branched and develop as prolongation of the lower

epidermal cells. They are of two types: (i) Smooth-walled rhizoids, (ii)
Tuberculate rhizoids.

In smooth-walled rhizoids both the inner and outer wall layers are fully stretched
while in tuberculate rhizoids appear like circular dots in surface view (Fig. 1 F).
The inner wall layer modifies into peg like in growth which projects into the cell
lumen (Fig. 1 H). The main functions of the rhizoids are to anchor the thallus on the
substratum and to absorb water and mineral nutrients from the soil.
Anatomy of the Gametophyte:
A vertical cross section of the thallus can be differentiated photosynthetic zone and
lower storage zone (Fig. 2 A, B, E).
Upper Photosynthetic zone:
The outermost layer is upper epidermis. Its cells are thin walled square, compactly
arranged and contain few chloroplasts. Its continuity is broken by the presence of
many barrel shaped air pores. Each pore is surrounded by four to eight
superimposed tiers of concentric rings.
(Fig. 2 B) with three to four cells in each tier (Fig. 2 D).

Air pores are compound in nature. The lower tier consists of four cells which
project in the pore and the opening of the pore looks star like in the surface view
(Fig. 2 C). The walls of the air pore lie half below and half above the upper
epidermis (Fig. 2 B).

Just below the upper epidermis photosynthetic chambers are present in a horizontal
layer (Fig. 2 B). Each air pore opens inside the air chamber and helps in exchange
of gases during photosynthesis.
These are chambers develop schizogenously (Vocalized separation of cells to form
a cavity) and are separated from each other by single layered partition walls. The
partition walls are two to four cells in height. Cells contain chloroplast. Many
simple or branched photosynthetic filaments arise from the base of the air chambers
(Fig. 2 B).

Storage zone:
It lies below the air chambers. It is more thickened in the centre and gradually
tapers towards the margins. It consists of several lasers of compactly arranged, thin
walled parenchymatous isodiametric cells. Intercellular spaces are absent.

The cells of this zone contain starch. Some cells contain a single large oil body or
filled with mucilage. The cells of the midrib region possess reticulate thickenings.
The lower most cell layer of the zone forms the lower epidermis. Some cells of the
middle layer of lower epidermis extend to form both types of scales and rhizoids
(Fig. 2 B).

Reproduction in Marchantia:
Marchantia reproduces by vegetative and sexual methods.
(i) Vegetative Reproduction:
In Marchantia it is quite common and takes place by the following methods:
1. By Gemmae:
Gemmae are produced in the gemma cups which are found on the dorsal surface of
the thallus (Fig. 3 A). Gemma cups are crescent shaped, 3 m.m. in diameter with
smooth, spiny or fimbriate margins (Fig. 3 B).

V. S. passing through the gemma cup shows that it is well differentiated into
two regions:
Upper photosynthetic region and inner storage region (Fig. 3 D).

The structure of both the zones is similar to that of the thallus. Mature gemmae are
found to be attached at the base of the gemma cup by a single celled stalk.

Intermingled with gemmae are many mucilage hairs. Each gemma is autotrophic,
multicellular, bilaterally symmetrical, thick in the centre and thin at the apex. It
consists parenchymatous cells, oil cells and rhizoidal cells. It is notched on two
sides in which lies the growing point (Fig. 3 C).
All cells of the gemma contain chloroplast except rhizoidal cells and oil cells.
Rhizoidal cells are colourless and large in size. Oil cells are present just within the
margins and contain oil bodies instead of chloroplast.

Dissemination of Gemmae:
Mucilage hairs secrete mucilage on absorption of water. It swells up and presses the
gemmae to get detached from the stalk in the gemma cup. They may also be
detached from the stalk due to the pressure exerted by the growth of the young
gemmae. The gemmae are dispersed over long distances by water currents.

Germination of Gemmae:
After falling on a suitable substratum gemmae germinate. The surface which comes
in contact with the soil becomes ventral surface.

The rhizoidai cells develop into rhizoids. Meanwhile, the growing points in which
lies the two lateral notches form thalli in opposite directions. Thus, from a single
gemmae two thalli are formed. Gemmae which develop on the male thalli form the
male plants and those on the female thalli form the female plant.

Development of Gemma:
The gemma develops from a single superficial cell. It develops on the floor of a
gemma cup. It is papillate and called gemma initial (Fig. 3 E). It divides by a
transverse division to form lower stalk cell and upper cell (Fig. 3 F). The lower cell
forms the single celled stalk.

The upper cell further divides by transverse division to form two cells. Both cells
undergo by similar divisions to form four cells (Fig. 3 G). These cells divide by
vertical and horizontal division to form a plate like structure with two marginal
notches. It is called gemma (Fig. 3 H-J).

2. Death and decay of the older portion of the thallus or fragmentation:

The thallus is dichotomously branched. The basal part of the thallus rots and
disintegrates due to ageing. When this process reaches up to the place of
dichotomy, the lobes of the thallus get separated. The detached lobes or fragments
develop into independent thalli by apical growth (Fig. 4 A-C).

3. By adventitious branches:
The adventitious branches develop from any part of the thallus or the ventral
surface of the thallus or rarely from the stalk and disc of the archegoniophore in
species like M. palmata (Kashyap, 1919). On being detached, these branches
develop into new thalli (Fig. 4 D).

(ii) Sexual Reproduction:

Sexual reproduction in Marchantia is oogamous. All species are dioecious. Male
reproductive bodies are known as antheridia and female as archegonia. Antheridia
and archegonia are produced an special, erect modified lateral branches of thallus
called antheridiophore and archegoniophore arpocephalum) respectively (Fig. 5 A,
B).

Further growth of the thallus is checked because growing point of the thallus is
utilised in the formation of these branches. In some thalli of M. palmatci and L.
polymorpha abnormal receptacle bearing both anheridia and archegonia have also
been reported, such bisexual receptacles are called as androgynous receptacles.
Internal structure of Antheridiophore or Archcgoniophore:
Its transverse section shows that can be differentiated into two sides: ventral side
and dorsal side. Ventral side has two longitudinal tows with scales and rhizoids.
These grooves, run longitudinally through the entire length of the stalk. Dorsal side
shows an internal differentiation of air chambers. (Fig. 5 C).

Antheridiophore:
It consists of 1-3 centimetre long stalk and a lobed disc at the apex (Fig. 32). The
disc is usually eight lobed but in M. geminata it is four lobed. The lobed disc is a
result of created dichotomies.

L.S. through disc of Antheridiophore:

The disc consists of air chambers alternating with heridial cavities. Air chambers
are more or less triangular and open on upper surface by n pore Called ostiole.
Antheridia arise in acropetal succession i.e., the older near the center and youngest
at the margins. (fig. 6 A).

Mature Antheridium:
A mature antheridium is globular in shape and can be differentiated into two parts
stalk and body. Stalk is short multicellular and attaches the body to the base of the
antheridial chamber. A single layered sterile jacket encloses the mass of androcyte
mother cells which metamorphosis into antherozoids (Fig. 6 B, 7 G). The
antherozoid is a minute rod like biflagellate structure (Fig. 8 H).
Development of Antheridium:
The development of the antheridium starts by a single superficial cell which is
situated on the dorsal surface of the disc, 2-3 cells behind the growing point. This
cell is called antheridial initial (Fig. 7 A). The antheridial initial increases in size
and divides by a transverse division to form an outer upper cell and a lower basal
cell (Fig. 7 B).

Basal cell remains embedded in the tissue of the thallus, undergoes a little further
development and forms the embedded portion Of the antheridial stalk. Outer cell
divides to form a filament of four cells. Upper two cells of the four celled filament
are known as primary antheridial cells and lower two cells are known as primary
stalk cells (Fig. 7 C).
Primary stalk cells from the stalk of the antheridium. Primary antheridial cells
divide by two successive vertical divisions at right angle to each other to form two
tiers of four cells each
(Fig. 7 D). A periclinal division is laid down in both the tiers of four cells and there
is formation of eight outer sterile jacket initials and eight inner primary androgonial
cells (Fig. 7 E).

Jacket initials divide by several anticlinal divisions to form a single layer of sterile
antheridial jacket. Primary androgonial cells divide by several repeated transverse
and vertical divisions resulting in the formation of large number of small
androgonial cells (Fig. 7 F).

The last generation of the androgonial cells is known as androcyte mother cells
(Fig. 7 G). Each androcyte mother cells divides by a diagonal mitotic division to
form two triangular cells called androcytes. Each androcyte cell metamorphosis
into an antheozoid (Fig. 8 A-G).
Spermatogenesis:
The process of metamorphosis of androcyte mother cells into antherozoids is called
spermatogenesis.

It is completed in two phases:

(1) Development of blepharoplast.

(2) Elongation of androcyte nucleus.

1. Development of Blepharoplasty:
In the young triangular androcyte (Fig. 8 D) blepharoplast appears as a dense
granule in one of the acute angles. It elongates to some extent and puts its whole
body in close contact with the inner contour of androcyte. From the elongated
blepharoplast emerge the flagella.

2 Elongation of Androcyte nucleus:

With the elongation of blepharoplast, the nucleus also elongates. The spline
apparatus acts as a cytoskeleton for the elongation of nucleus. Spline apparatus is a
multilayered structure which comprises tubules (Fig. 8 E-H).

Archegoniophore or Carpocephalum:
It arises at the apical notch and consists of a stalk and terminal disc. It is slightly
longer than the antheridiophore. It may be five to seven cm. long. The young apex
of the archegoniophore divides by three successive dichotomies to form eight lobed
rosette like disc.
Each lobe of the disc contains a growing point. The archegonia begin to develop in
each lobe in acropetal succession, i.e., the oldest archegonium near the centre and
the young archegonium near the apex of the disc. (Fig. 10 A). Thus, eight groups of
archegonia develop on the upper surface of the disc. There are twelve to fourteen
archegonia in a single row in each lobe of the disc.

Development:
The development of the archegonium starts on the dorsal surface of the young
receptacle in acropetal succession. A single superficial cell which acts as
archegonial initial enlarges and divides by transverse division to form a basal cell
or primary stalk cell and an outer cell or primary archegonial cell (Fig. 9 A, B).

The primary stalk cell undergoes irregular divisions and forms the stalk of the
archegonium.
The primary archegonial cell divides by three successive intercalary walls or
periclinal vertical walls resulting in the formation of three peripheral initials and a
fourth median cells, the primary axial cell (Fig. 9 C, D).
Each of the three peripheral initials divide by an anticlinal vertical division forming
two cells (Fig. 9 G, H) In this way primary axial cell gets surrounded by six cells.
These are called jacket initials (Fig. 9 H, I). Six jacket initials divide transversely
into upper neck initials and lower venter initials (Fig. 9 F). Neck initial tier divides
by repeated transverse divisions, to form a tube like neck.

Diversity of Algae, Lichens & Bryophytes:

Neck of the archegonium consists of six vertical rows. (Fig. 9 I). Each row consists
six to nine cells Venter initials tier also divides by rapid transverse divisions to
form a single wall layer of swollen venter (Fig 9 K). Simultaneously, the primary
axial cell divides transversely and unequally to form upper small primary cover cell
and lower large central cell (Fig. 9 E).

The central cell divides into primary neck canal cell and a lower venter cell.
Primary neck canal cells divides by a series of transverse divisions to form a row of
about eight thin walled neck canal cells (Fig. 9 J, K).

Primary venter cell divides only once and forms a small upper venter canal cell and
a lower large egg or ovum (Fig. 9 K). The primary cover cell divide by two vertical
divisions at right angle to one another to form four cover cells which form the
mouth of the archegonium.

Mature Archegonium:
A mature archegonium is a flask shaped structure. It remains attached to the
archegonial disc by a short stalk. It consists upper elongated slender neck and basal
globular portion called venter. The neck consists of six vertical rows enclosing
eight neck canal cells and large egg.
Four cover cells are present at the top of the neck. (Fig. 9 L).

Fertilization in Marchantia:
Marchantia is dioecious. Fertilization takes place when male and female thalli grow
near each other. Water is essential for fertilization. The neck of the archegonium is
directed upwards on the dorsal surface of the disc of the archegoniophore (Fig. 9
A).

In the mature archegonium the venter canal cell and neck canal cells disintegrate
and form a mucilaginous mass. It absorbs water, swells up and comes out of the
archegonial mouth by pushing the cover cells apart. This mucilaginous mass
consists of chemical substances.
The antherozoids are splashed by rain drops. They may fall on the nearby female
receptacle or swim the whole way by female receptacle. It is only possible if both
the male and female receptacles are surrounded by water.

Many antherozoids enter the archegonial neck by chemotactic response and reach
up to egg. This mechanism of fertilization is called splash cup mechanism. One of
the antherozoids penetrates the egg and fertilization is effected. The fusion of both
male and female nuclei results in the formation of diploid zygote or oospore.
Fertilization ends the gametophytic phase.

Sporophytic Phase:
Post Fertilization Changes:
After Fertilization the following changes occur simultaneously:
1. Stalk of the archegoniophore elongates.

2. Remarkable over-growth takes place in the central part of the disc. As a result of
this growth the marginal region of the disc bearing archegonia is pushed
downward and inward. The archegonia are now hanging towards the lower side
with their neck pointing downwards (Fig. 10 B-D).

3. Wall of the venter divides to form two to three layered calyptra.

4. A ring of cells at the base of venter divides and re-divides to form a one cell
thick collar around archegonium called perigynium (Pseudoperianth).

5. A one celled thick, fringed sheath develops on both sides of the archegonial row.
It is called perichaetium or involucre. Thus, the developing sporophyte is
surrounded by three protective layers of gametophytic origin i.e., calyptra,
 perigynium and perichaetium (Fig. 11). The main function of these layers is to
provide protection, against drought, to young sporophyte.

6. Between the groups of archegonia, long, cylindrical processes develop from the
periphery of disc. These are called rays. They radiate outward, curve downwards
and give the disc a stellate form. In M. polymorpha these are nine in number.

7. Zygote develops into sporogonium.

Development of Sporogonium:
After fertilization the diploid zygote or oospore enlarges and it completely fills the
cavity of the archegonium. It first divides by transverse division (at right angle to
the archegonium axis) to form an outer epibasal cell and inner hypo basal cell (Fig.
12 A, B).
The second division is at right angle to the first and results in the formation of four
cells. This represents the quadrant stage (Fig. 12 C). The epibasal cell forms the
capsule and hypo basal cells form the foot and seta.

Since the capsule is developed from the epibasal cell and forms the apex of the
sporogonium, the type of embryogeny is known as exoscopic. The next division is
also vertical and it results in formation of eight celled stage or octant stage.

Now the divisions are irregular and globular embryo is formed (Fig. 12 D). The
lower cells divide to form a massive and bulbous foot. The cells of the seta divide
in one plane to form vertical rows of cells.

In upper region of capsule (when the young sporogonium is about a dozen or more
cells in circumference) periclinal division occurs and it differentiates it into outer
single layered amphithecium and multilayered endothecium (Fig. 12 E, F).

The cells of the endothecium divide only by anticlinal divisions to form a single
layered sterile jacket or capsule wall. The endothecium forms the archesporium. Its
cells divide and re-divide to form a mass of sporogenous cells (sporocytes). Half of
the sporogenous cells become narrow and elongate to form the elater mother cells.
(Fig. 12 G, I).

In M. polymorpha sporogenous cells divide by five successive divisions to form

thirty-two spore mother cells while in M. domingensis sporogenous cells divide
only by three to four divisions to form eight or sixteen spore mother cells. The
elater mother cells elongate considerably to form long, slender diploid cells called
elaters.

Elaters are pointed at both the ends and have two spiral bands or thickenings on the
surface of the wall. These are hygroscopic in nature and help in dispersal of spores
(Fig. 12 K). The spore mother cell is diploid and divides meiotically to form four
haploid spores which remain arranged tetrahedrally for quite some time (Fig. 12 J).
The spores later become free and remain enclosed by the capsule wall along elaters.
(Fig. 12 H).
The quadrant type of development of sporogonium is quite common in many
species of Marchantia (e.g., M. polymorpha) but in a few species zygote divides by
two transverse divisions to form the 3-celled filamentous embryo. In it the hypo
basal cell forms the foot, the middle seta and the epibasal cell develops into
capsule. However, it is the rare type of embryo development in M. chenopoda.

Mature Sporogonium:
A mature sporogonium can be differentiated into three parts, viz., the foot, seta and
capsule (Fig. 13 H). Foot. It is bulbous and multicellular. It is composed of
parenchymatous cells. It acts as anchoring and absorbing organ. It absorbs the food
from the adjoining gametophytic cells for the developing sporophyte.

Seta:
It connects the foot and the capsule. At maturity, due to many transverse divisions it
elongates and pushes the capsule through three protective layers viz., calyptra,
perigynium and perichaetium.

Capsule:
It is oval in shape and has a single layered wall which encloses spores and elaters. It
has been estimated that as many as 3, 00,000 spores may be produced in single
sporogonium and there are 128 spores in relation to one elater.

Dispersal of Spores:
As the sporogonium matures, seta elongates rapidly and pushes the capsule in the
air through the protective layers (Fig. 13 A). The ripe capsule wall dehisces from
apex to middle by four to six irregular teeth or valves. The annular thickening in the
cells of the capsule wall causes the valves to roll backward exposing the spores and
elaters.

The elaters are hygroscopic in nature. In dry weather they lose water and become
twisted. When the atmosphere is wet, they become untwisted and cause the jerking
action. Due to this the spore mass loosens and spores are carried out by air currents
(Fig. 13 B, C).

Structure of Spore:
Spores are very small (0.012 to 0.30 mm in diameter). They are haploid,
uninucleate, globose and surrounded by only two wall layers. The outer well layer
is thick, smooth or reticulate and is known as exospore or exine. The inner wall
layer is thin and is called endospore or intine. In M. torsana and M. caneiloba they
are tetrahedrally arranged.

Germination of Spores and Development of Gametophyte:

Under favourable conditions, the spores germinate immediately. In first year the
spore viability is approximately 100%. Before germination it divides by transverse
division to form two unequal cells (Fig. 14 A, B). The lower cell is small in size.

It is relatively poor in cell contents, achlorophyllous and extends to form germ-

rhizoid (Fig. 14 C). The large cell is chlorophyllous and undergoes divisions to
form a six to eight cell germ-filament or protonema (Fig. 14 D). At this stage the
contents of the cells migrate at the apex.

The apex is cut off from the rest of the sporeling by a division. It behaves as apical
cell. It is wedge-shaped with two cutting faces. The apical cell cuts off five to seven
cells alternately to the left and right. These cells by repeated divisions form a plate
like structure (Fig. 14 F).

According to O’ Hanlon’s (1976) a marginal row of cells appears in the apical

region in this plate. By the activity of these marginal cells, the expansion of the
plate takes place into thallus, a characteristic of Marchantia.

Marchantia is dioecious, 50% of the spores develop into male thalli and 50%
develop into female thalli (Fig. 15).
Alternation of Generation in Marchantia:
The life cycle of Marchantia shows regular alternation of two morphologically
distinct phases. One of the generations is Haplophase and the other is diplophase.

(i) Haplophase or Gametophytic Phase:

In Marchantia this phase is dominant and produces the sex organs. Sex organs
produce gametes to form a diploid zygote.

(ii) Diploid Phase or Sporophytic Phase:

Zygote develops into sporophyte. In Marchantia sporophyte is represented by foot,
seta and capsule. The sporophyte produces the spores in the capsule. The spores on
germination produce the gametophyte.

So, in Marchantia two morphologically distinct phases (Haplophase and

Diplophase) constitute the life cycle. The life cycle of this type which is
characterised by alternation of generations and sporogenic meiosis is known as
heteromorphic and diplohaplontic (Fig. 16).