Lycopodium: Habit and Habitat

and Morphology
In this article we will discuss about:- 1. Habit and Habitat of
Lycopodium 2. External Morphology of Lycopodium
3. Internal Structure 4. Reproduction 5. Morphological
Nature of Protocorm 6. Economic Importance 7. Life Cycle
Patterns.
Habit and Habitat of Lycopodium:
Lycopodium is commonly known as ‘club moss’ due to their moss like
appearance and club shaped strobili. It has about 400 species, which
are cosmopolitan in distribution. They are found in colder arctic
region as well as in temperate, tropical and sub-tropical regions but
they are abundantly found in tropical zones.

Thirty three species of Lycopodium have been reported from India.

Mostly it is found growing in moist and shady places which are rich in
humus and other organic matters. Some of the common species are L.
clavatum, L. phlegmaria, L. cernuum, etc.

It has got 2 sub-genuses:

(i) Urostachya—branching dichotomous and roots originate from the
base of the stem.

(ii) Rhopalostachya—stem prostrate with erect branching and roots

arise adventitiously from all along the stem.

Mostly the tropical species are epiphytic (e.g., L. phlegmaria) and grow
hanging from the tree trunks. The temperate species may be erect and
shruby (e.g., L. reflexum), creeping (e.g., L clavatum) or erect form
(e.g., L. cernuum) etc.
External Morphology of Lycopodium:
The herbaceous plant body is sporophytic. Usually they may have
either prostrate stem with erect leafy branches or weak pendent stem
(epiphytes).

The plant body is distinctly differentiated into following

three regions (Fig. 1 A-C):
(i) Stem,

(ii) Roots, and

(iii) Leaves.

(i) Stem:
In the sub-genus Urostachya stem is erect (terrestrial) or pendent
(epiphytic) and may be branched (dichotomously) or unbranched. In
the sub-genus Rhopalostachya the stem is prostrate with erect
branches. First the branching is dichotomous and later on becomes
monopodial.

(ii) Root:

Usually small, adventitious roots are present. In the sub-genus

Urostachya roots originate only from the base of the stem (not arising
from the whole length of the stem). In some species e.g., L. selago etc.
the roots arise endogenously from pericycle of the stem, do not
penetrate the cortex of the stem but turn downward through the cortex
and finally emerge only at the base of the stem.

Due to this reason a T. S. of stem usually shows roots within the cortex
and are known as cortical roots (inner roots). In sub-genus
Rhopalostachya also roots are adventitious and arise all along the
underside of the prostrate portion of the stem.
(iii) Leaves:
Leaves are simple, sessile, small in size, eligulate and possess a single
unbranched midrib and are known as microphylls. Usually the leaves
are spirally arranged (e.g., L. clavatum) but may be arranged in whorls
(e.g., L. cernuum) or pairs (e.g., L. alpinum).

In all the cases they condensely cover the surface of the stem. Leaves
are usually homophyllous (isophyllous) i.e., of same size and shape but
in some cases e.g., in L. complanatum the leaves are heterophyllous
(anisophyllous) i.e., of different size.

Usually the leaves near the apical portion of the branches bear
sporangia and are called sporophylls. Depending upon the species the
sporophylls may or may not be differentiated from the ordinary leaves.

These sporophylls usually form a condense structure at the apex of the

branches which are known as strobili. The numbers of strobili at the
tip of branches differ in different species.
Internal Structure of Lycopodium:
(a) Stem:
A transverse section (T.S.) of the stem of Lycopodium is
somewhat circular in outline and can be differentiated into
following three regions:
1. Epidermis:
It is the outermost covering layer comprising of single cell in
thickness. The epidermis is cutinised on the outer side and interrupted
at places by the presence of stomata.

2. Cortex:
Inner to the epidermis is present a wide zone of cortex which shows a
great variation in its structure in different species.
Usually four types of cortex are recognized:
(i) The whole of the cortex is made up of parenchymatous cells with
small or large intercellular spaces (e.g., L. selago). Such cortex is called
homogeneous.

(ii) The whole of the cortex is made up of sclerenchymatous cells,

without intercellular spaces.

(iii) The cortex is differentiated into outer and inner sclerenchymatous

cells and middle parenchymatous cells (e.g., L. clavatum, Fig. 2 A).

(iv) The cortex is differentiated into outer and inner parenchymatous

cells and middle sclerenchymatous cells (e.g., L. cernuum Fig. 2. B).
Next to the cortex is present a single layer of well-defined cells known
as endodermis with conspicuous casparian strips but at maturity the
endodermis may or may not be a distinct structure. Endodermis is
followed by pericycle which is composed of one or more layers of
compactly arranged parenchymatous cells.

3. Stele:
It is made up of only primary xylem and primary phloem. It is a
protostele i.e., pith is absent and the stele is situated in the centre. The
arrangement of xylem and phloem tissues is different in different
species and the stele is also named differently.

In case of L. serratum, L. phlegmaria etc. the xylem is star shaped with

a protoxylem situated at the periphery (protoxylem exarch Fig. 3 A). In
L. annotinum in actinostele the furrows in the xylem are much more
and show stellate arrangement (Fig. 3B).

The phloem lays in the space between the xylem rays. This type of stele
is known as actinostele. In case of L. clavatum. L. volubile etc. xylem
appears to be in the form of separate plates arranged somewhat
parallel, with phloem in between them.
This type of stele is known as plectostele (Fig. 2 A, 3 C). In case of L.
cernuum, L. drummondii etc. xylem and phloem are uniformly
distributed i.e. it appears as if strands of xylem are embedded in the
phloem. This type of stele is known as mixed protostele (Fig. 2 B, 3 D).

The protoxylem is usually exarch in all the cases. Xylem is usually

composed of tracheids and phloem of sieve tubes and phloem
parenchyma. Cambium is absent hence there is no secondary growth
i.e., no formation of secondary xylem and secondary phloem.

(b) Root:
The roots are adventitious.
A transverse section (T.S.) of the aerial root of Lycopodium is
somewhat circular in outline and shows the following internal
structure:

(i) Epidermis:
It is the outermost covering layer and is only one cell thick. The cells
are thin walled. Epidermis is provided with numerous root hairs
present in pairs (characteristic of Lycopodium).

(ii) Cortex:
Just below the epidermis is present a wide zone of cortex. It is
differentiated into outer sclerenchyma and inner parenchyma. The
outer one gives the mechanical strength to the root.
(iii) Stele:
It may be di-, tetra-, or polyarch. In prostrate species it is polyarch i.e.,
having 6-10 plates of xylem arranged radially (star shaped). The xylem
is exarch. The phloem is present between the radiating arms of xylem.
In erect or pendent species stele is diarch or tetrarch. In L.selago, L.
serratum it is diarch and xylem is C, U or crescent shaped. The phloem
is present between the 2 ends of xylem, only in one group.
The cortical roots are exactly similar in their internal structure to that
of aerial roots, except that the epidermis and root hairs are absent.

The xylem is composed of tracheid and phloem of sieve tubes and

phloem parenchyma. The endodermis and pericycle are indistinct
structure at maturity. Pith and cambium are absent.

(c) Leaf:
T. S. of the leaf shows epidermis, mesophyll tissue and a
single median vascular bundle:
1. Epidermis:
It is the outermost surrounding layer and is only one cell in thickness.
The cells of epidermis are parenchymatous and cutinised on their
outer side. The epidermis is also interrupted by the presence of
stomata. In homophyllous (isophyllous) species the stomata are
present on outer as well as inner epidermis (amphistomatic) but in
heterophyllous (anisophyllous) species the stomata are mostly
restricted on the lower epidermis (hypostomatic).
2. Mesophyll:
It occupies a wide zone between the epidermis and the vascular
bundle. It is usually made up of thin walled chlorenchymatous cells
which may be with or without intercellular spaces.

3. Vascular bundle:
In the centre of the leaf is situated only a single concentric vascular
bundle made up of only xylem and phloem. The vascular bundle is
surrounded on all sides by a sclerenchymatous sheath.

Reproduction in Lycopodium:
Lycopodium reproduces by two methods vegetatively and by spores.

1. Vegetative reproduction:
It takes place by the following methods:
(i) Gemmae or bulbils:
In a few species like L. selago, L. lucidulum etc. certain buds like
structures known as gemmae or bulbils are usually produced in large
number on new stem tips annually. The morphological nature of
gemmae is still not fully known. The gemmae when fall on ground,
develop root primodia and soon form the root.
(ii) Death and decay:
Species with creeping stem reproduces vegetatively by the death and
decay of older parts of the stem up to the point of branching. This
separates the branches which later on grow independently.
(iii) Resting buds:
In L. inundatum the whole of the plant body except the growing tip of
rhizome is dead during winter. This tip portion of the rhizome acts as
resting bud which in the coming spring resumes growth and develops
into a new plant.
(iv) Fragmentation:
In several epiphytic species fragments of the plant body are capable of
giving rise to new plants.
2. Sexual Reproduction:
Spore Producing Organs:
Lycopodium is a sporophytic plant and reproduces sexually. The
plants are homosporous i.e., produces only one type of spores (without
differentiation of mega- and microspores). These spores are produced
in sporangia which, in turn, are produced on fertile leaves known as
sporophylls. Usually the sporophylls are grouped together to form a
compact structure known as strobili (Sing. strobilus) which are
terminal structures (Fig. 1 A).

Strobilus (Reproductive organ):

In the primitive species of the sub-genus Urostachya every leaf on the
plant is a sporophyll or at least potentially so and the fertile and sterile
zones alternate. The sporophylls are loosely arranged. In species of
Rhopalostachya and in some species of Urostachya the leaves of the
apical portion of the branches only bear sporangia and are called
sporophylls. The rest behave as vegetative leaves.

The sporophylls may be of the same size or of different size from the
foliage leaves in different species. The arrangement of sporophylls is
same on the central axis as that of the vegetative leaves on the stem
i.e., spiral, whorled or decussate etc.

The position of the sporangium is also different in different species.

The sporangia may be axillary and protected with the help of
sporophylls (e.g., L. inundatum Fig. 7 A) or foliar and protected (e.g.,
L. cernuum Fig. 7 B) or subfoliar and exposed (e.g., L. squarrosum,
Fig. 7 C) or axillary and exposed (e.g., L. lucidulum, Fig. 7 D).

Longitudinal section (L.S.) of strobilus shows the presence of a

strobilus axis in the centre. On both sides of the strobilus axis are
present sporophylls (Fig. 8 A). Each sporophyll bears only one
sporangium (Fig. 8 B). All the sporangia are similar in structure and
are arranged acropetally in a strobilus i.e., the youngest are at the top
(Fig. 8 C).
Structure of Sporangium:
Sporangia are sac-like structures but usually kidney shaped in
appearance (Fig. 8 B). Sometimes they are sub-spherical in
appearance. Their colour varies from orange to yellow. Each
sporangium consists of a basal short massive stalk i.e., sub-sessile,
with an upper globular unilocular body containing numerous spores.

The body of the sporangium consists of 3 or more layers of wall

surrounding a cavity. The inner most layer of the wall of sporangium is
called as tapetum (Fig. 9 F) which is nutritive in nature and persists
till maturity.

In the young sporangium inside the wall is present a mass of

sporogenous cells which in due course of development form spore
mother cells which by meiotic divisions, produce haploid tetrad of
spores. The spores at maturity separate from each other.
The wall of the sporangium is provided with a transverse strip of cells
known as stomium from where the sporangium at maturity splits into
2 valves and the spores are dispersed away in the air.

The spores produced by a sporangium are all alike (homosporous).

They are small, rounded or even spherical structures. The surface of
the spores is usually rough due to the presence of reticulate ridges or
knob like protrusions. Each spore is provided with a triradiate ridge
(Fig. 8, D, E) and is somewhat yellow in colour. A small amount of
chlorophyll may or may not be present in spores. Reserve food is in
the form of oil in the spores.

Development of sporangium and formation of spores. Bower (1894)

had studied the development of sporangium in Lycopodium. The
sporangium develops from a small group of superficial cells arranged
in a transverse row on the adaxial side of the sporophyll near the base.

Its development is of eusporangiate type. These superficial cells are

called sporangial initials (Fig. 9A, B). These cells divide by periclinal
divisions forming an outer and inner layer of cells. The outer cells
divide periclinally and anticlinally forming three celled thick wall of
the sporangium (Fig. 9A-F).

The inner layer or archesporial cells divide in all directions forming a

group of cells known as sporogenous tissue which finally give rise to
spore mother cells. During these developments the inner-most layer of
wall is differentiated as a nutritive layer and is known as tapetum. It is
a persistent structure and rich in reserve food material.

Each spore mother cell undergoes a process of meiosis thus producing

a tetrad of spores (haploid) with tetrahedral arrangement. These
spores later on separate from the tetrad, as a result of which, a large
number of spores are produced inside each mature sporangium.
Dehiscence of sporangium and liberation of spores. As the sporangium
approaches towards maturity, a transverse row of cells is
differentiated near the apical portion from the wall of a sporangium
known as stomium.

The walls of the cell of stomium thicken and differ from the walls of
other cells of the sporangium. As the sporangium loses water, it
creates a pressure on the wall which leads to the appearance of slit in
the stomium as a result of which the wall splits opens into two halves
and the spores are disseminated by air current.

Gametophytic Generation:
The development of the gametophyte (prothallus) takes place from the
haploid spores which are the unit of gametophytic generation. Each
spore is unicellular, uninucleate haploid structure, 0.03 mm in
diameter and surrounded by 2 layers, with a triradiate ridge at the
surface (Fig. 8 D, E).

Chlorophyll may or may not be present in different species. In few

species spores may germinate within a few days after liberation but in
some species the spores germinate when they are 3-8 years old and the
development of gametophyte upto formation of mature sex organs
may take a time of 8 months to 6 or even 15 years.
The rate of the formation of photosynthetic tissue is usually
proportional to the rate of growth of gametophyte. Both the male and
female reproductive organs are produced on the same gametophyte.
The male sex organs are produced earlier than female sex organs.

Usually at the time of germination of spore, it swells up to absorb

water. First the spore divides into two unequal cells by a lenticular
division, forming a very small lens shaped cell known as rhizoidal cell
and a bigger cell (Fig. 10 A, B).

This rhizoidal cell takes no part in further development of

gametophyte and is a colourless structure. At this two celled stage the
spore will rupture at the triradiate ridge. Second division divides the
bigger cell into two equal halves, the cell near the rhizoidal cell is
known as basal cell and the other one is known as upper cell (Fig. 10
C).

The upper cell further divides by two successive divisions in such a

way as to form an apical cell with two cutting faces (Fig. 10 D). At this
stage the gametophyte is 5 celled structures and the symbiotic
phycomycetous fungus (mycorrhizal fungus) attacks it.

If this fungus fails to attack at this stage, further development of

gametophyte stops. This infection takes place through the basal cell.
During further course of development of gametophyte the apical cell
further divides to form six or morecells which later on develop into
meristematic cells. These cells, by further divisions form a
multicellular structure, the gametophyte (prothallus) (Fig 10 E-H).
Structure of the Mature Gametophytes:
The form and structure of the gametophytes varies greatly in different
species.

In general they have been grouped under three categories:

Type I or Cernuum type:
Gametophyte is partially aerial and partly in soil. The prothalli are
usually 2 to 3 millimetre in height and 1-2 millimetre in diameter. The
gametophytes (prothalli) grow at the surface of the ground and consist
of a colourless basal portion buried in soil and a conspicuous upright,
fleshy, green aerial portion having lobes (Fig. 11 A).
The sex organs develop between the green expanding lobes. The
prothallus itself is a nourishing body. The underground part contains
endophytic fungus e.g., L. cernuum, L. inundatum etc.

Type II or Clavatum Type:

The gametophyte is wholly subterranean and totally saprophytic i.e.,
non- green structure. It is tuberous and without lobes. It may be one to
two centimentre long or wide and is top shaped, conical or discoid in
shape (Fig. 11 B, C). The endophytic fungus is present. Sex organs are
formed on the upper surface e.g. L. annotinum, L. complanatum, L.
clavatum etc.

Type III or Phlegmaria type:

The gametophyte is subterranean, saprophytic and colourless. This
type of prothallus is seen in L. phlegmaria and other epiphytic species.
The prothallus is about 2 millimeter in diameter and monopodially
branched (Fig. 11 D). Sex organs are borne on upper surface of large
branches and are interspersed with slender filaments known as
paraphyses.
Besides these three forms some intermediate forms of prothalli are
also observed. In L. selago the prothalli may be subterranean or
epiterranean (aerial). If the spores are buried under the soil after
liberation, they form subterranean prothalli and if the spores are not
buried under soil after their liberation, they form epiterranean
prothalli.
The internal structure of the prothallus is very simple. The outermost
layer is epidermis, followed by cortical mycorrhizal region, palisade
region and central storage region. It is attached with the substratum
by unicellular rhizoids. The prothalli of all species are monoecious i.e.,
antheridia and archegonia develop on the same prothallus.

Development of sex organs:

Both the sex organs i.e., antheridia (male) and archegonia (female)
develop on the same prothallus, usually in distinct patches on the
upper surface. The gametophytes are protandrous i.e., antheridia
develop before archegonia. Sex organs develop just on the back of the
apical meristem.
Development of antheridium:
A single superficial cell situated just away from the meristematic cells
gives rise to an antheridium. This superficial cell is known as
antheridial initial (Fig. 12 A). This cell divides periclinally to form an
outer cell known as jacket initial (primary wall cell) and an inner cell
known as primary androgonial initial or cell (Fig. 12 B).

The jacket initial divides only anticlinally by several divisions resulting

in the formation of single layered covering known as jacket layer. In
the middle of the jacket layer a triangular cell is differentiated, which
is known as opercular cell.

Simultaneously, the primary androgonial divides by various divisions,

forming a mass of cells embedded in the prothallus, known as
androgonial cells which finally give rise to androcytes (antherozoid
mother cells, Fig. 12 C-F). The number of androcytes per antheridium
varies in different species.

Each androcyte later on metamorphosis into a biflagellated

antherozoid. Each antherozoid is a haploid, uninucleate, fusiform
structure with broad rounded posterior end and an upper pointed
biflagellated anterior end (Fig- 12 G).

The triangular opercular cell becomes mucilaginous as a result of

which an opening is formed at the apex of antheridium through which
water enters into it. The antherozoids absorb water and swell up as a
result of which a pressure is created on the wall of antheridium which
finally ruptures and the antherozoids are liberated.

Development of archegonium:
Just like antheridium, the archegonium also arises from a single
superficial cell called archegonial initial, situated just away from the
meristematic cells at the apex (Fig. 13 A). The archegonial initial
divides by periclinal division into an upper primary cover cell and
lower basal central cell (Fig. 13 B).

The primary cover cell later on divides vertically by two successive

divisions at right angle to each other forming four neck initials which
later on by transverse divisions form a 3-4 cells high neck. Each tier of
the neck consists of 4 cells.
The central cell divides transversely forming an, upper primary canal
cell and a lower primary ventral cell (Fig. 13 D). The primary canal cell
by successive transverse divisions produces a variable number of neck
canal cells (usually one in L. cernuum, seven in, L. selago and 14-16 in
L. complanatum).

The primary ventral cell may directly behave as an egg or may divide
transversely to form an upper ventral canal cell and a lower egg (Fig.
13 E-G). The egg is somewhat broader then the rest part of
archegonium. The archegonial jacket is absent. The archegonium is a
sunken flask shaped structure with neck projecting out of the
prothallus.
Fertilization:
At the time of fertilization the neck canal cells and the ventral canal
cell disorganise and the cells of the upper-most tier of neck slightly
separate apart forming a passage upto the egg (Fig. 13 H). Fertilization
is brought about in the presence of water.

The biflagellate antherozoids reach the archegonium by swimming in

water on the surface of prothallus. The antherozoids are perhaps
attracted towards the neck of archegonium by a chemotactic
movement. They enter the archegonium through neck and reach the
egg.

Only the nucleus of one antherozoid fuses with the egg nucleus thus
forming a diploid structure-known as oospore (2x). The act of
fertilization ends the gametophytic generation and the initial stage of
sporophytic generation is formed.
Embryo Development (Young Sporophyte):
The rate of development of the embryo is extremely slow. In
Lycopodium embryo develops downward into the gametophytic tissue
instead of developing upward i.e., towards the neck of archegonium.
The first division of the oospore is always transverse, forming an
upper cell (epibasal) and a lower cell (hypobasal) known as embryonic
cell.

The upper cell does not divide further and behaves as suspensor. The
lower cell (embryonic cell) divides by two vertical divisions at right
angle to each other, followed by a transverse division, forming 8 cells
(octant, Fig. 14 A-D). The 4 cells of the octant, situated near the
suspensor by further division, form a multicellular foot which acts as a
haustorium and helps in the absorption of food material from the
gametophytic tissue.

Out of the 4 remaining cells of the octant, the 2 cells towards the
meristematic region give rise to stem and the other 2 cells give rise to
primary leaf and primary root (Fig. 14 D-J). The primary stem is short
lived and is replaced by adventitious outgrowth which gives rise to
horizontal stem. More roots develop from the stem.

The primary roots of the sporophyte are exogenous in origin while

those arising later on are endogenous in origin. The embryo obtains its
nourishment for a long time from the gametophyte.

In some species e.g., L. cernuum etc. the gametophyte is generally

green. The oospore normally divides transversely forming suspensor
and embryonic cell. The embryonic cell forms an octant. The tier
which gives rise to stem, leaf and primary roots, develops into a
massive spherical structure of parenchymatous cells, known as
protocorm (Fig. 14 K, L).

It grows through the gametophyte, becomes green and develops

rhizoids on its lower surface. The upper surface of the protocorm gives
rise to a few to many erect outgrowths which are leaf like and are
known as protophylls.

The protophylls are provided with stomata. At this stage the

protocorm separates from the gametophyte. Now at the upper side of
protocorm a region is differentiated which develops into stem.
Protocorm is regarded as the intermediate phase in between normal
embryo and definite leafy shoot.
Morphological Nature of Protocorm of Lycopodium:
Various views have been put forward to explain the morphological
nature of protocorm of Lycopodium.

A few important ones are discussed below:

(1) Treub (1837) regarded the protocrorm as the remains of primitive
undifferentiated structure originally possessed by the Pteridophytes
and in majority of the present day Pteridophytes it has been replaced
by a definite leafy shoot. This view is now only of historical
importance.

(2) Bower (1908, 1935) regarded it as a swelling of occasional

adaptation. It acts as an organ of perennation. It has no phylogenetic
importance.

(3) Holloway (1910) regarded it as a specialised structure that helps

the young sporophyte to perennate over dry season.
(4) Browne (1913) regarded it as a modified and a reduced stem.

(5) Wardlaw (1955) regarded it as a modified shoot.

Economic Importance of Lycopodium:

Different species of Lycopodium are differently important as for
example, some species of Lycopodium (L. obscurum) are used in
making Christmas wreaths. L. volubile is used for table decoration.

Extract from the plant of Lycopodium was used as kidney stimulant in

the old times. The spores of Lycopodium are highly inflammable and
have been used to produce stage lighting, for the theatres. The spores
of L. clavatum etc. are used in pharmacy as water repellent protective
dusting powder for tender skin etc.

Life Cycle Patterns in Lycopodium:

Lycopodium is a sporophyte (2x) with distinct sporophytic (2x) and
gametophytic (x) generations which alternate with each other. The
plant is homosporous i.e., reproduces by producing only one type of
spores. The spore on germination produces gametophyte (x) which, in
turn, produces both antherozoids and eggs in antheridia and
archegonia respectively.

These reproductive structures later on after fertilization produce

zygote (2x) which again on germination gives rise to a sporophytic
plant (2x). In this way sporophytic and gametophytic generations
alternate with each other and it shows a distinct alternation of
generation although the sporophytic phase is dominant over
gametophytic phase (Fig. 15).
Equisetum: Habitat, Structure
and Reproduction
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In this article we will discuss about:- 1. Habit and Habitat of

Equisetum 2. Structure of Equisetum 3. Reproduction 4. Life
Cycle.
Habit and Habitat of Equisetum:
The plant body of Equisetum has an aerial part and an underground
rhizome part (Fig. 7.83). The rhizome is perennial, horizontal,
branched and creeping in nature. The aerial part is herbaceous and
usually annual. Majority of the species are small with a size range in
between 15 and 60 cm in height and 2.0 cm in diameter.

However some species grow up in higher heights [e.g., E. giganteum

(13 m), E. telmateia (2 m); E. ramosissimum (4 m), though their stem
are relatively thin (0.5-2.0 cm in diam.)] showing vine-like habit and
climb over adjacent forest trees.
Equisetums generally grow in wet or damp habitats and are
particularly common along the banks of streams or irrigation canals
(E. debile, E. palustre). However, some species are adapted to xeric
condition (e.g., Equisetum arvense). Some common Indian species are
: E. arvense, E. debile, E. diffusum, E. ramosissimum.

Some species of Equisetum are indicators of the mineral content of the

soil in which they grow. Some species accumulate gold (about 4.5
ounce per ton of dry wt.), thus they are considered as ‘gold indicator
plants.

Hence these plants help in prospection/exploration for new ore

deposits. In Equisetum, silica is deposited on the outer wall of the
epidermal cells giving the characteristic rough feeling, thus it provides
a protective covering against predators and pathogens.

Structure of Equisetum:
The Sporophyte:
ADVERTISEMENTS:
The sporophytic plant body of Equisetum is differentiated into stem,
roots and leaves (Fig. 7.83).

Stem:
The stem of Equisetum has two parts: perennial, underground, much-
branched rhizome and an erect, usually annual aerial shoot. The
branching is monopodial, shoots are differentiated into nodes and
internodes.

In majority of the species, all the shoots are alike and chlorophyllous
and some of them bear strobili at their apices (e.g., E. ramosissimum,
E. debile). Sometimes shoot shows dimorphism (two types of shoots
i.e., vegetative and fertile) e.g., E. arvense.

Some shoots are profusely branched, green (chlorophyllous) and

purely vegetative. The others are fertile, unbranched, brownish in
colour (achlorophyllous) and have terminal strobili.

The underground rhizome and the aerial axis appear to be articulated

or jointed due to the presence of distinct nodes and internodes.
Externally, the internodes have longitudinal ridges and furrows and,
internally, they are hollow, tube-like structures. The ridges of the
successive internodes alternate with each other and the leaves are
normally of the same number as the ridges on the stem.

Internal Features of Stem:

In T.S., the stem of Equisetum appears wavy in outline with ridges and
furrows (Fig. 7.84). The epidermal cell walls are thick, cuticularised
and have a deposition of siliceous material.
Stomata are distributed only in the furrows between the ridges. A
hypodermal sclerenchymatous zone is present below each ridge which
may extend up to stele in E. giganteum. The cortex is differentiated
into outer and inner regions.

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The outer cortex is chlorenchymatous, while the inner cortex is made

up of thin-walled parenchymatous cells. There is a large air cavity in
the inner cortex corresponding to each furrow and alternating with the
ridges, known as vallecular canal. These are schizolysigenous canals
extending the entire length of internodes and form a distinct aerating
system.

New leaves and branches of Equisetum are produced by the apical

meristem, however, most of the length of the stem are due to the
activity of intercalary meristem located just above each node. The
activity of intercalary meristem causes rapid elongation of the inter-
nodal region.
The stele is ectophloic siphonestele which is surrounded by an outer
endodermal layer. An inner endodermis is also present in some
species of Equisetum (e.g., E. sylvaticum). The endodermis is followed
by a single-layered pericycle.

The vascular bundles are arranged in a ring which lies opposite to the
ridges in position and alternate with the vallecular canals of the cortex.
Vascular bundles are conjoint, collateral and closed. In the mature
vascular bundle, protoxylum is disorganised to form a carinal cavity
which lies opposite to the ridges.

The metaxylem tracheids (scalariform or reticulate) are present on

both sides of the phloem. In some species vessels with reticulate
perforations are reported. The central part of the internode of aerial
shoot is occupied by a large pith cavity which is formed due to rapid
elongation of the internodal region.

The vascular bundles remain unbranched until they reach the level of
node. At the nodal region, each vascular bandle trifurcates (divided
into three parts).

The middle branch of the trifurcation enters the leaf. Each lateral
branch of the trifurcate bundle joins a lateral strand of an adjacent
trifurcate bundle to form a vascuiar bundle of internode (Fig. 7.85).
Thus the vascular bundles of internode alternate with those of inter-
nodes above and below.

In the nodal region, the xylem is extensively developed as a

conspicuous circular ring. There are no vallecular or carinal canals at
this level. In addition, a plate of pith tissue occurs at the node which
separates one internode from another.
The internal structures of the shoot of Equisetum is peculiar because it
shows xerophytic as well as hydrophytic features.

The xerophytic features are:

(i) Ridges and furrows in the stem,

(ii) Deposition of silica in the epidermal cells,

(iii) Sunken stomata,

(iv) sclerenchymatous hypodermis,

(v) Reduced and scaly leaves, and

(vi) photosynthetic tissue in the stem.

The hydrophytic characteristics on the other hand are (i) we 11-

developed aerating system like carinal canal, vallecular canal and
central pith cavity, and (ii) reduced vascular elements.

Root:
The primary root is ephemeral. The slender adventitious roots arise
endogenously at the nodes of the stems. In T.S., the root shows
epidermis, cortex and stele from periphery to the centre. The
epidermis consists of elongated cells, with or without root hairs.
The cortex is extensive; cells of the outer cortex often have thick walls
(sclerenchymatous) and those of the inner cortex are thinner
parenchymatous. The stele is protostelic where the xylem is triarch or
tetrarch, or, in smaller roots, may be diarch.

A large metaxylem element is present in the centre of the stele and the
protoxylem strands lie around it. The space between the protoxylem
groups is filled with phloem. There is no pith.

Leaves:
The leaves of Equisetum are small, simple, scale-like and isophyllous;
they are attached at each node, united at least for a part of the length
and thus form a sheath around the stem. The sheath has free and
pointed teeth-like tips.

The number of leaves per node varies according to the species. The
species with narrow stems have few leaves (e.g., 2-3 leaves in E.
scirpoides) and those with thick stem have many leaves (e.g., up to 40
leaves in E. schaffneri).

The number of leaves at a node corresponds to the number of ridges

on the internode below. The leaves do not perform any photosynthetic
function and their main function is to provide protection to young
buds at the node.

Reproduction in Equisetum:
Equisetum reproduces vegetatively and by means of spores.

i. Vegetative Reproduction:
The subterranean rhizomes of some species (e.g., E. telmateia, E.
arvense) form tubers (Fig. 7.83) which, on separation from the parent
plant, germinate to produce new sporophytic plants. The tubers
develop due to irregular growth of some buds at the nodes of the
rhizomes.

ii. Reproduction by Spores:

Spores are produced within the sporangia. The sporangia are borne on
the sporangiophores which are aggregated into a compact structure
termed strobilus or cone or sporangiferous spike.

Strobilus:
The strobilus are terminal in position and generally are borne
terminally on the chlorophyllous vegetative shoot (Fig. 7.86A).
However, they may be borne terminally on a strictly non-
chlorophyllous axis (e.g., E. arvense).

The strobilus is composed of an axis with whorls of sporangiophores

(Fig. 7.86B, C). Each sporangiophore is a stalked structure bearing a
hexagonal peltate disc at its distal end (Fig. 7.86D). On the under
surface of the sporangiophore disc 5-10 elongate, cylindrical hanging
sporangia are borne near the periphery in a ring.

The flattened tips of the sporangiophores fit closely together which

provide protection to the developing sporangia. The axis bears a ring-
like outgrowth, the so-called annulus immediately below the whorls of
sporangiophores which provide additional protection during early
development.

The annulus has been interpreted as a rudimentary leaf sheath by

some botanists, whereas others consider it to be sporangiophoric in
nature as occasionally it bears small sporangia.

Development of Sporangium:
The mode of development of sporangium is eusporangiate, as it is not
entirely formed from a single initial. Superficial cells adjacent to the
original initial may also take part in the development of sporangium.

Sporangia are initiated in single superficial cell around the rim of the
young sporangiophore. The periclvnal division of the sporangium
initial forms an inner and an outer cell. The inner cell, by further
divisions in various planes, gives rise to sporogenous tissue.
The outer cell, by periclinal and anticlinal divisions, gives rise to
irregular tiers of cells, the inner tiers of which may transform into
sporogenous tissue and the outer tiers become the future sporangial
wall cells.

The innermost layer of the sporangial wall differentiates as the

tapetum. The sporogenous cells separate from each other, round off
and eventually transform into spore mother cell. All but the two
outermost wall layers disorganise to form periplasmodial fluid.

However, not all of the sporogenous cells function as spore mother

cells. Many of them degenerates to form a multinucleate nourishing
substance for the spore mother cells. Each spore mother cell
undergoes meiotic division (reductional division) and produces spore
tetrad. All spores in a sporangium are of same size and shape i.e.,
homosporous.

Structure of Mature Sporangium:

The mature sporangium is an elongated saclike structure, attached to
the inner side of the peltate disc of the sporangiophore (Fig. 7.86D). It
is surrounded by a jacket layer which is composed of two layers of
cells. The inner layer is generally compressed and the cells of the outer
layer have helical thickenings which are involved in sporangial
dehiscence.

Dehiscence of Sporangium:
At maturity, the strobilar axis elongates, as a result the
sporangiophores become separated and exposed. Then the
sporangium splits open by a longitudinal line due to the differential
hygroscopic response of the wall cells.

Spores:
The spores are spherical and filled with densely packed chloroplasts.
The spore wall is laminated and shows four concentrate layers. The
innermost is the delicate intine, followed by thick exine, the middle
cuticular layer and the outermost epispore or perispore. The intine
(endospore) and exine (exospore) are the true walls of the spore.
The outer two layers i.e., cuticular layer and epispore are derived due
to the disintegration of the nonfunctional spore mother cells and
tapetal cells. At maturity, the epispore (the outermost layer) splits to
produce four ribbon like bands or strips with flat spoon-like tips.

These bands are free from the spore wall except for a common point of
attachment and remain tightly coiled around the spore wall until the
sporangium is fully matured.

These are called elaters (Fig. 7.87A). The elaters are hygroscopic in
nature. The spores remain moist at early stages of development, thus
the elaters are spirally coiled round the spore. The spores dry out at
maturity and consequently the elaters become uncoiled.

These uncoiled elaters become entangled with the elaters of other

spores. Through these actions the elaters help in the dehiscence
process and also the dispersal of spores in large groups from the spo-
rangium.

The elaters of Equisetum are different from those of the bryophytes

(Table 7.6).

Gametophyte Generation:
Equisetum is a homosporous pteridophyte. The haploid spores
germinate to form gametophyte. The germination takes place
immediately if the spores land on a suitable substratum. If the spores
do not germinate immediately, their viability decrease significantly.

The spores swell up by absorbing water and shed their exine (Fig.
7.87B). The first division of the spore results in two unequal cells: a
small and a large cell (Fig. 7.87C). The smaller cell elongates and
forms the first rhizoid. The larger cell divides irregularly to produce
the prothallus. The prevailing environmental conditions determine the
size and shape of the prothallus.

If a large number of spores are developed together within a limited

space, then the prothalli formed are of thin filamentous type. But a
relatively thick and cushion-shaped prothalli are formed from sparsely
germinating spores. Mature gametophytic plants may range in size
from a few millimeters up to 3 centimeters e.g., E. debile) in diameter.

They are dorsiventral and consist of a basal non-chlorophyllous

cushion-like portion from which a number of erect chlorophyllous
muticellular lobes develop upwards. Unicellular rhizoids are formed
from the basal cells of cushion (Fig. 7.87D).

The prothallus bears sex organs and reproduces by means of sexual

method.

Sexuality in Equisetum:
The gametophytic plant body bears sex organs i.e., antheridium (male)
and archegonium (female). The gametophyte are basically bisexual
(homothallic) i.e., they bear both male and female sex organs (Fig.
7.87D). Although, some unisexual (dioecious) members are also
reported (Fig. 7.87E, F). Some are initially unisexual and then become
bisexual.

This early sex determination appears to be related to the

environmental conditions viz., temperature, light, humidity and the
supply of nutrients as well. Ducket (1977), in order to explore the
sexuality in Equisetum, observed that some of the fragments of male
gametophyte remained male throughout the successive subcultures
under laboratory conditions.

Some other fragments produced archegonia, which subsequently bore

antheridia in increasing numbers. This phenomenon supports the
contention that Equisetum gametophytes are potentially bisexual.
However, Hanke (1969) observed that gametophyte of Equisetum
bogotense were unisexual (bearing antheridia) and never change to
bisexual type.

However, the initial male gametophyte of E. ramosissimum, E.

variegatum and E. bogotense never became bisexual.

Schratz (1928) observed that 50% spores germinate to produce male

gametophytes, while the remaining 50% spores produce female
gametophytes though they do not loose their male potentiality (i.e.,
antheridia develop later if fertilisation fails). He termed this as
‘incipient heterospory’.

A study of sexuality based on enzymatic analysis revealed the

intragametophytic self- fertilisation in E. arvense.

Equisetum is homosporous and, therefore, definite sex-determining

mechanism is absent. But, the sexuality demonstrated by some of the
members appears to be related to environmental factors. Therefore, it
is termed as environmental sex determination.
Sex Organs of Equisetum:
i. Antheridium:
In monoecious species, antheridia develop later than archegonia. They
are of two types — projecting type and embedded type. Antheridia first
appear on the lobes of the gametophyte (Fig. 7.87D). The periclinal
division of the superficial antheridial initial gives rise to jacket initial
and an androgonial cell (Fig. 7.88A, B).

The jacket initial divides anticlinally to form a single-layered jacket.

The repeated divisions of androgonial cells form numerous cells
which, on metamorphosis, produce spermatids/antherozoids (Fig.
7.88C-E). The antherozoids escape through a pore created by the
separation of the apical jacket cell.

The apical part of the antherozoid is spirally coiled, whereas the lower
part is, to some extent, expanded (Fig. 7.88F). Each antherozoid has
about 120 flagella attached to the anterior end.

ii. Archegonium:
Any superficial cell in the marginal meristem acts as an archegonial
initial which undergoes periclinal division to form a primary cover cell
and an inner central cell (Fig. 7.89A, B). The cover cell, by two vertical
divisions at right angle to each other, forms a neck (Fig. 7.89C). The
central cell divides transversely to form a primary neck canal cell and a
venter cell (Fig. 7.89D).

Two neck canal cells are produced from the primary neck canal cell.
While, the venter cell, by a transverse division, forms the ventral canal
cell and an egg (Fig. 7.89E).

At maturity, an archegonium has a projecting neck comprising of three

to four tiers of neck cells arranged in four rows, two neck canal cells of
unequal size, a ventral canal cell, and an egg at the base of the embed-
ded venter (Fig. 7.89F-G). The archegonia are confined to cushion
region in- between the aerial lobes (Fig. 7.87D).
Fertilisation:
Water is essential for fertilisation. The gametophyte must be covered
with a thin layer of water in which the motile antherozoides swim to
the archegonia. The neck canal cells and ventral canal cell of the
archegonia disintegrate to form a passage for the entry of
antherozoids.
Many antherozoids pass through the canal of the archegonium but
only one of them fuses with the egg. Thus diploid zygote is formed.
Generally more than one archegonia are fertilised in a prothallus.

Embryo (The New Sporophyte):

The embryo is the mother cell of the next sporophytic generation.
Unlike most pteridophytes, several sporophytes develop on the same
prothallus. The first division of the zygote is transverse. This results in
an upper epibasal cell and lower hypobasal cell. The embryo is there-
fore exoscopic (where the apical cell is duacted outward i.e., towards
the neck of the archegonium) in polarity.

No suspensor is formed in Equisetum. The epibasal and hypobasal

cells then divide at right angles to the oogonial wall, and as a result a
tour-celled quadrant stage is established (Fig. 7.90A). All the four cells
of the quadrant are of different size and shape.

The four-celled embryo undergoes subsequent divisions and the future

shoot apex originate from the largest cell and leaf initials from the
remaining cells of one quadrant of the epibasal hemisphere.

One cell of the epibasal quadrant and a portion of the adjacent

quadrant of the hypobasal region contribute to the development of
root. The first root develops from one of the epibasal quadrants and a
portion of the adjacent hypobasal quadrant. The shoot grows rapidly.
Later the root grows directly downward and penetrate the
gametophytic tissue to reach the soil or substratum (Fig. 7.90B, C). A
number of such sporophytes may develop from a large mature
gametophyte if more than one egg is fertilised (Fig. 7.90D).

Life Cycle of Equisetum:

Fig. 7.91 depicts the life cycle of Equisetum.
Adiantum: Occurrence and
Gametophyte | Botany
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In this article we will discuss about:- 1. Occurrence and Distribution of

Adiantum 2. Sporophyte of Adiantum 3. Gametophyte 4. Phylogeny.

Occurrence and Distribution of Adiantum:

Adiantum is popularly called ‘Maiden hair fem’ because of the shiny
black rachis of the leaves. It is one of the most widely distributed
genera (Other genera are Cheilanthes, Pellaea, Ceratopieris and
Anogramma) of the family growing luxuriantly in both tropical and
sub topical regions of the world. It grows ubiquitously wherever nature
offers a moist, shaded locality. There are nearly 200 species.

Nayar (1961) has investigated the morphology of 24 Indian species of

Adiantum. Some of the common Indian species are – A. capillus-
veneris, A.pedatum, A. incisum, A. caudatum, A. venustum, A.
lunulatum, A. edgowrthii etc. Species of Adiantum are commonly
cultivated in green houses because of their attractive foliage.

Sporophyte of Adiantum:
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Morphology of the plant: The sporophytic plant body consists of an

underground rhizome from which are produced leaves and roots. The
rhizome is covered with chaffy scales (Paleae). It may be erect (A.
caudatum), semi erect (A. pet- datum), or creeping (A.capillus-
veneris).

Studies of Nicholas (1985) in Adiantum trapeziforme indicate that the

erect rhizome of the young sporophyte quickly transforms itself into
creeping. The rhizome may be hard or soft and brown in colour.

The chaffy scales that cover the rhizome are of various shapes and
sizes. Nayar (1961) has made a detailed study of these scales in 24
species of Adiantum. From the undersurface of the rhizome arise a
number of adventitious roots. The roots are stiff and black in colour.
Occasionally they may be branched.

The leaves are produced in acropetalous succession on the creeping

rhizome. They show circinate vernation typical of ferns. The rachis of
the leaf is hard, wiry, shiny and black or dark brown in colour thus
giving the name maiden hair fern. The rachis has a medium dorsal
groove, and is covered with paleae at the basal region. In addition to
this, glandular hairs may also be present.

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The leaves may be unipinnate (Axaudatum) or bi or tri-pinnate as in ,

A. capillus – veneris (Fig. 149). The pinnae are stalked and have a
dichotomous venation. The rachis may terminate in a pinna or may
bear a bud. In A capillus veneris the rachis divides pinnately and the
ultimate branches bear pinnae in an alternate fashion.

There is no distinction between fertile and sterile leaves in Adiantum.

The whole leaf may be sporangiferous or only certain pinnae may bear
sporangia. The soral organisation is very evident. Sori are borne on the
ventral surface of the pinnae.
Apical Organization:
In a study of apical organisation of rhizome, leaf and root in Adiantum
capillus veneris, Bir and Randhawa (1984) have reported the
occurrence of a single apical cell, which is later replaced by a group of
cells.

Internal Structure:
1. Rhizome:
A transection reveals the usual three zones epidermis, cortex and stele
(Fig. 150). The outline of the section would be wavy. Epidermis is
single layered and the cells may be thin walled or thick walled. There is
a cuticle external to the epidermis.

Cortex lies internal to the epidermis. It may be wholly

parenchymatous (A. rubellum), (Fig. 150) or it may have sclerenchyma
and parenchyma. In A. pectinatum, scattered masses of sclerenchyma
are found embedded in the parenchymatous ground tissue. In A.
caudatum, sclerenchyma constitutes the hypodermal region.

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The central vascular cylinder exhibits great variety. In A. capillus

veneris, it is a dictyostele consisting of a ring of meristeles. In the
young condition the stele may be a solenostele. In A. rubellum the
stele is a typical amphiphloic solenostele, with characteristic features
such as outer endodermis, outer pericycle, outer phloem, xylem, inner
phloem, inner pericycle and inner endodermis lining the
parenchymatous pith.
2. Leaf:
The petiole shows an epidermis, parenchymatous cortex and the vas-
cular trace. Srivastava (1979) has studied the foliar epidermis of
Adiantum. There is a thick walled hypodermis next to the epidermis.
The number of leaf traces entering the leaf, varies.
It is single in A. caudatum and others and double in A. capillus –
veneris. Even when there are two leaf traces, both of them unite
further up resulting in a single bundle. The xylem is concave at the
base but triradiate higher up with three protoxylem groups. Xylem is
exarch.

In A. bausei there is a patch of included parenchyma in the xylem.

Khare and Shankar (1986) studied the vascular organisation of the
petiole in Adiantum caudatum, A.edgeworthii, A.pedatum,
A.phillippense, A.pubescens and A.trapeziforme and have reported
two types of vascular supply to the leaf.

In A.phillipense, A.caudatum and A.edgeworthii there is always a

single vascular trace from the rhizome which remains unaltered in the
petiole, while in the other three species two distinct traces originate
from the rhizome.

These two merge into one after entering the petiole. In a comparative
anatomical study of the stipe of Adiantum, Bidin and Walker (1985)
have reported eight different types of xylem configurations. According
to them this is of systematic value.

The lamina shows the two epidermal layers upper and lower the
mesophyll is generally undifferentiated. It is highly reduced in A.
capillus – veneris, A pedatum, etc., having only two layers of cells. In
A. pedatum, in some regions the mesophyll is totally absent and at
such places the two epidermal layers are closely appressed to each
other.

The mesophyll (when present) as well as the epidermal layers are

chlorophyllous. The epidermal layers are chlorophyllous. The
epidermal cells over the veins are thick walled. The stomata are
scattered throughout the surface of the leaf. Paleae or ramenta may be
borne even on the epidermis of the lamina. The vein may or may not
have a bundle sheath. The vascular tissues show the characteristic X\P
arrangement.
3. Root:
A transection shows a very prominent piliferous layer, a two zoned
cortex and the central protostele (Fig. 152). The piliferous layer has
brown coloured cell walls. Cortex has an outer parenchymatous zone
and an inner sclerotic zone.

Surrounding the stele is a conspicuous endodermis with prominent

casparian thickenings. The xylem is exarch and diarch, phloem
completely surrounds the xylem. External to phloem is a single layered
pericycle.
Reproduction:
Vegetative propagation is brought about by buds produced at the leaf
tips. The buds enter the ground when the leaf bends and touches the
soil. There they develop into a new individual. This, in turn may repeat
the process leading to the walking Habit. Walking habit is seen in A
caudatum.

Spore Producing Organs:

As has already been said there is no distinction into fertile and sterile
leaves. The son are born at the distal end of the pinnae. But the sori
are not exactly marginal. They are borne a little behind the tip of the
veins.

The sorus bearing margin of the leaf incurls and forms the false
inducium. In some cases sporangia may develop at the distal ends of
the veins (A. phillippense). In the sori paraphyses may be present in
between the sporangia as in A. rubellum, A. tenerum, etc. The sorus is
of the mixed type.
Development and Structure of the Sporangium:
The development is similar to what is seen in Pteris. A mature
sporangiuim has a stalk made up of three rows of cells. The stalk
terminates in a globose or biconvex capsule. The wall is single layered.

There is an obliquely vertical annulus (Fig.153) of 12-24 cells long. The

annulus is separated from the stalk by two or three cells. The stomium
also is separated from both the stalk, and the annulus. The rest of the
sporangial wall is composed of a few large cells.

The sporangium dehisces transversely liberating the spores. All the

spores are of the same type.

Gametophyte of Adiantum:
Structure and germination of the spores:
Spores are tetrahedral in shape. The wall is two layered. Exine is thick
and smooth and has a brownish tinge. On falling upon a suitable
substratum the spore germinates. The first sign of germination is the
rupturing of exine and the protruding out of the germ tube.

The germ tube undergoes several transverse divisions to form a short

filament. The lowest cell (Fig. 154a) forms a lateral rhizoid. The
terminal cell becomes an apical cell with three cutting faces. By the
division of the apical cell, a spatulate pro-thallus is formed first. (Fig.
154).

The mature pro-thallus is cordate, photosynthetic, dorsiventrally

flattened and aerial. The growing point is situated in the apical notch
(Fig.154e, 154f). All the cells in the pro-thallus are parenchymatous.
The pro-thallus is one celled thick towards the margins but many
celled thick towards the centre. In some species collenchyma may be
found at the corners. Rhizoids are produced from the ventral surface.

Reproduction:
The prothalli are monoecious. Antheridia are found in between the
rhizoids towards the ventral surface. Arehegonia are found near the
growing point towards the ventral surface. Structure and development
of sex organs is same as in Pteris.

Embryogeny:
The first division of the zygote is vertical (Fig. 155b). The epibasal half
(next to the archegonial neck) forms the leaf and root while the hypo
basal half forms the stem’ apex and foot (Fig. 155c). Embryogeny is
essentially similar to what is seen in Pteris.

Generally only one sporophyte is formed per pro-thallus. During

embryogeny the root and juvenile leaves make their appearance first,
with the stem differentiating late. The primary root penetrates the soil
and establishes itself. Apogamy has been reported in A.philippense.

Chromosome Number:
In a cytological study of 51 species of ferns from western India,
Mahabale and Kamble (1981) have reported polyploidy in Adiantum.
In another cytological study of the genus (A.peruvianum) Sinha and
Verma (1984) have reported a chromosome number of 2n = 60, with
an irregular meosis.

Phylogeny of Adiantum:
Adiantaceae seems to occupy a none too high position among the
mixtae. The lack of a true inducium is one of the characteristic features
of the family. Considering the relationships of polypodiaceae, Eames
(1964) opines that gymnogrammoids (to which Adiantum belongs) are
themselves polyphyletic and are probably associated with
osmundaceous and schizaeaceous stock.
Marsilea: Habit, Structure and
Reproduction
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In this article we will discuss about:- 1. Habit of Marsilea

2. Structure of Marsilea 3. Reproduction.
Habit of Marsilea:
The plants are herbaceous with rhizomatous stem, creeping on or just
below the soil surface.
The species of Marsilea are generally aquatic or amphibious in nature
with their roots embedded in mud or damp soil. The aquatic species —
M. minuta and M. quadrifolia — occur in fresh water ponds, shallow
water, or in mud or damp soil.
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When grown in water, the whole body is submerged with just the leaf
lamina floating on the surface of water. The amphibious species, M.
aegyptiaca occur in seasonally wet habitats which produces repro-
ductive structures during the drier periods.
M. condensata and M. rajasthanensis are near xerophytic forms which
grow in dry soil. M. hirsuta is an extremely xerophytic plant reported
from Australia.
Structure of Marsilea:
Sporophyte:
The sporophytic plant body of Marsilea shows differentiation of stem,
leaves and roots (Fig. 7.116).
1. Stem:
The stem is long, slender and freely-branched rhizome of indefinite
growth that grows on or just below the soil surface. It is differentiated
into nodes and internodes. The internodes are generally long in
aquatic species but are short in sub- terrestrial or terrestrial species.
The production of underground tubers on rhizome has been reported
only in M. hirsuta.
A T.S. of the rhizome (stem) shows epidermis, cortex and the stele
from periphery to the centre (Fig. 7.11 7).
The outermost layer is epidermis, composed of compactly arranged
thick-walled cells and are devoid of stomata. The cortex is extensive
and differentiated into three layers viz., outer cortex, middle cortex,
and inner cortex. The outer cortex is parenchymatous with large air
spaces.
The air chambers are separated from each other by a single-layered
septum. The middle cortex is sclerenchymatous, while the inner cortex
is made up of compactly arranged parenchymatous cells. The air
spaces are absent in xerophytic species (e.g., M. aegyptiaca).
The stele is amphiphloic solenostelic which occupies the centre of the
rhizome. Xylem occurs in the form of a ring and is surrounded on
either side by phloem.
The central part of the stele is occupied by pith which is
parenchymatous in aquatic species and sclerenchymatous in
xerophytic species (e.g., M. aegyptiaca). The stele is bounded
externally by outer pericycle and outer endodermis, while it is
bounded internally by inner pericycle and inner endodermis.
2. Leaves:
ADVERTISEMENTS:
The leaves arise from the nodes and are arranged alternately in two
rows on the upper side of the creeping rhizome. The leaves are long
petiolate and palmately compound, each having four leaflets in many
species, but sometimes the number of leaflets varies from 3-8. A young
leaf shows circinate vernation. At maturity the pinnae are extended
perpendicular to the petiole. The venation is of closed reticulate type.
In T.S., the petiole differentiates into epidermis, cortex and stele (Fig.
7.118). The epidermis is cutinised and composed of a single-layered
rectangular cells.
The cortex is differentiated into outer and inner cortex. The outer
cortex consists of aerenchyma having many air-cavities separated by
one-celled thick septa.
The stele is protostelic with diarch and exarch xylem. The xylem has
two large metaxylem elements at the centre and protoxylem elements
at each end towards the periphery. Phloem bands are present on either
side of Fig. 7.118 : Marsilea minuta. T.S. of petiole the xylem.

The stele is protostelic, generally triangular in outline and is externally

bounded by endodermis. A single-layered pericycle is present just
inside the endodermis.
The xylem is mesarch, consists of two outwardly curved strands.
Protoxylem patches are present at both the ends with one or more
metaxylem in the centre. The xylem is surrounded by phloem. Pith is
absent.
The vertical section of the leaflet shows epidermis, stomata, mesophyll
and vascular bundles. Stomata are present in the upper epidermis (in
submerged species) or on both the upper and lower epidermis (in
terrestrial and amphibious species).
Mesophyll tissue is differentiated into palisade and spongy
parenchyma and generally associated with vertical airspaces. The
vascular bundles are concentric with a central core of xylem
surrounded by phloem. Each vascular bundle is separated from the
mesophyll tissue by a layer of endodermis.
3. Root:
The primary roots are short-lived (ephemeral) and are replaced by
adventitious roots. Roots usually arise at nodes on the lower side of
the rhizome. However, in some species (e.g., M. aegyptiaca) the roots
arise from the internodal region of the rhizome.
In transverse section, the root shows three distinct regions : the
epidermis, the cortex, and the stele (Fig. 7.119).
The epidermis is composed of single-layered parenchymatous cells,
outer walls of which are thickly cuticularised.
The cortex is differentiated into three zones, viz., the outer cortex, the
middle cortex, and inner cortex. The outer cortex is aerenchymatous
with many air spaces separated by radially elongated septa. The
middle cortex is parenchymatous. The inner cortex is thick-walled,
sclerenchymatous, internally bounded by the successive layers of
endodermis and pericycle.
Reproduction in Marsilea:
Marsilea reproduces vegetatively as well as by means of the spores.
i. Vegetative Reproduction:
Under some unfavourable circumstances the subterranean branches of
the rhizome form tubers. These structures have reserve food in the
form of oil globules which help them to overcome the unfavourable
conditions. On return of the favourable conditions, these tubers germi-
nate and form new plant body (e.g., M. hirsuta, M. minuta, M. erosa).
ii. Reproduction by Spores:
Marsilea is a heterosporous fern. It produces two types of spores i.e.,
the microspores and the megaspores. The microspores and
megaspores are produced in microsporangia and megasporangia,
respectively, and the sporangia are enclosed in special bean-shaped
structures called sporocarps.
When young, the sporocarps are soft and green, but turns dark brown
and hard at maturity. Sporocarps withstand desiccation and are repor-
ted to be viable even after twenty to twenty five years.
The sporocarp develops at the short branch of petiole called pedicel or
stalk. In most species they occur singly, but in some species the num-
ber varies from two to twenty (Fig. 7.120).
The mode of attachment of sporocarps with the petiole is of
following types:
(a) Sporocarps are attached to one side of the petiole in a single row
e.g., M. polycarpa, M. caribaca, M. subangulata, M. detlexa.
(b) Pedicels of the sporocarps are fused in groups and then attached to
the petiole by a common stalk e.g., M. quadrifolia.
(c) Pedicels of all the sporocarps remain free and are attached to the
petiole at a single point (e.g., M. minuta).
(d) A single sporocarp attached to the base of the petiole by the pedicel
(e.g., M. coro- mandelica, M. uncinata).

The place of attachment of the pedicel with the body of the sporocarp
is known as raphe (Fig. 7.121 A). The distal end of the raphe in some
species is marked by the presence of one or two protuberances or
teeth-like projections known as horns or tubercles.
The sporocarp wall is hard, thick, thus resistant against mechanical
injury. Anatomically, the wall is differentiated into three layers. The
outer layer is epidermis made up of single-layered cuboidal cell with
sunken stomata.
The middle layer is made up of radially elongated compactly arranged
thick-walled palisade cells. This is followed by second palisade layer
which is comprised of more elongated thin-walled palisade cells.
A vertical longitudinal section (VLS) in the plane of the stalk shows
that a single strong vascular strand enters the sporocarp near the
lower horn (Fig. 7.121B) and continues forward all along the dorsal or
upper side of the sporocarp thus forming a midrib (dorsal bundle).
From this midrib, the lateral side branches (lateral or commissural
bundles) arise which eventually pass on to the two sides of the
sporocarp. The sporocarp, therefore, has a bivalved structure.
Another bundle called placental bundle develops from the point of
forking of lateral bundle which enters into the receptacle bearing spo-
rangia and dichotomises. Thus a closed network of vascular system is
formed within the sporocarp.
A vertical longitudinal section (VLS) of sporocarp away from the plane
of the stalk reveals many sori arranged in vertical rows (Fig. 7.122). In
this plane of section either megasporangia or microsporangia are
visible. Each sorus is surrounded by an indusium. The development of
sori is of gradate type. The gelatinous mucilage ring is more prominent
in dorsal side.
A horizontal longitudinal section (HLS) cuts each sorus transversely
and it is seen that each sorus is an elongated structure, covered by a
delicate indusium.
The sori are gradate, basipetal in arrangement with a row of largest
sporangia (megasporangia) at top and two rows of smaller sporangia
(microsporangia) on two sides (Fig. 7.123). The mucilage ring is
present in the form of two masses, one in the dorsal and the other in
the ventral sides.

A vertical transverse section (VTS) of the sporocarp shows only the

sori on two sides (Fig. 7.124). Both the sori contain megasporangia if
the section is taken through the megasporangia or the sori contain
only microsporangia if it is taken through the microsporangia. The
sporophore is seen in the form of two masses on either side. The
mucilage ring is present only on the dorsal side.
Development of Sporangium:
Sporangial development is of the leptosporangiate type. The
development of both the micro- and megasporangia is almost alike.
The sporangial initials for megasporangium and microsporangium are
formed at top and at the sides of the receptacle, respectively.
The initial cell divides periclinally (transversely) into an outer and an
inner cell. The inner cell doe^ not take part in further development.
The outer cell undergoes three successive diagonal divisions and forms
a tetrahedral apical cell with three cutting faces.
This apical cell cuts off two segments along each of its three cutting
faces which forms the stalk of the sporangium. Now the apical cell
divides with the help of an arched periclinal wall towards its outer face
and forms an outer jacket initial and an inner tetrahedral archesporial
cell. The outer jacket initial divides anticlinally to form a single-
layered jacket.
The archesporial cell divides periclinally to forms an outer tapetal
initial and an inner primary sporogenous cell. Anticlinal and periclinal
divisions of the tapetal initial forms a two-layered thick tapetum.
The primary sporogenous cell divides to form a mass of either 8 or 16
spore mother cells. Spore mother cells (2n) undergo meiotic division
to form 32 or 64 haploid spores (n). The developments of both the
sporangia are similar up to this stage.
In megasporangium, only one megaspore survives to become a large
functional megaspore, while all the microspores are functional in
microsporangium.
Opening of the Sporocarp:
The sporangium wall of Marsilea shows no sign of cellular
specialisation (e.g., formation of annulus) required for dehiscence of
sporangium. A sporocarp is a hard structure and it does not open until
two or three years after their formation. This delay is probably due to
the imperviousness of the hard sporocarp wall.
Hence the sporocarp may remain viable for even 50 years. The tissues
slowly swell up by absorbing water in natural conditions. Thus the
swelling puts pressure on the wall of the sporocarp and eventually it
splits open along its ventral side into two halves.
Splitting is followed by the emergence of a long, worm-like gelatinous
structure to which the sori are attached (Fig. 7.125). The mucilaginous
cord may became ten or fifteen times larger than the sporocarp.
Following the release of the sori from the sporocarp, the indusia and
the sporangial wall disintegrate and the spores are liberated.

Gametophyte:
Marsilea is heterosporous i.e., they produce microspores and
megaspores which eventually germinate to form the male and female
gametophytes, respectively.
Male Gametophyte:
The microspores are small, globose structures with a thick outer
ornamented exine and inner thin intine. The outer exine is covered by
a thin layer called perispore. The microspore contains a distinct
haploid nucleus and its cytoplasm is rich in starch grains.
The microspores germinate inside the spore wall (endosporic type)
almost immediately after its release (Fig. 7.126A-G). It divides
asymmetrically to form a small prothallial cell and a large apical cell
(1-1). A division (2-2) of apical cell diagonal to prothallial cell forms
two antheridial cells.
Then both the antheridial cells divide diagonally (3-3) with curving
wall forming the first jacket cell and large wedge-shaped cell. The
jacket cells do not divide, but the wedge-shaped cell divides
periclinally (4-4) to form smaller inner cell (2nd jacket) and a large
outer cell. Further, the periclinal division (5-5) of outer cell forms 3rd
jacket and primary androgonial cell.
At this stage the male gametophyte consists of one prothallial cell, 6
jacket cells and 2 androgonial cells. After several divisions of the
primary androgonial cells, sixteen androcytes are formed surrounded
by jacket cells.
Later the prothalial cell and the jacket cells disintegrate and the two
groups of androcytes, representing the two antheridia, float freely in
the cytoplasmic mass within the original spore wall. Each androcyte
becomes a motile antherozoid by dissolution of the androcyte
membrane.
The antherozoids are corkscrew-shaped, multiflagellate structure cha-
racterised by the presence of a large posterior cytoplasmic vesicle (Fig.
7.126H).
Female Gametophyte:
The megaspore is an oval or elliptic structure, the wall of which
imbibes water and expands to form a gelatinous mass around the
megaspore. The spore wall expands to form a small papilla
(protuberance) at the apical end where the nucleus is located in a
dense part of cytoplasm. The remaining portion of the spore is filled
with a frothy cytoplasm full of starch grains (Fig. 7.127A).

The first division in the apical nucleus of the large megaspore is

transverse, forming a small nipple-shaped apical cell and a very large
basal nutritive or prothallial cell (Fig. 7.127B). The prothallial cell
provides the nutrition to the growing female gametophyte.
The apical cell further divides by three intersecting vertical walls to
establish an axial cell surrounded by three lateral cells (Fig. 7.127C).
Now the axial cell functions as archegonial initial which divides
periclinally to form an outer primary cover cell and an inner central
cell.
A small neck (2 tiers of 4 cells each) is derived from the primary cover
cell. The central cell divides transversely to form an upper primary
canal cell (behaves as neck canal cell) and lower primary venter cell
(Fig. 7.127D, E).
The primary ventral cell again divides transversely to form a ventral
canal cell and an egg. The growth of the archegonial complex ruptures
the megaspore wall at the apical end and forms a conspicuous
gelatinous mass with funnel-shaped papilla or protuberance. Now, the
megaspore splits through the triradiate fissure and archegonium
becomes exposed.
Fertilisation disintegration of the neck canal cell and ventral canal cell
create a passage for the antherozoids to fertilise the egg. The
antherozoids, after liberation from ruptured male gametophyte, enter
through the gelatinous protuberances and moves downwards to the
archegonium. One of the antherozoids eventually fuses with the egg to
form a diploid zygote.
New Sporophyte (The Embryo):
The zygote is the mother cell of the next sporophytic generation. The
first division of the zygote is vertical (in relation to the neck of the
archegonium) followed by a transverse division resulting in the
quadrant stage (four-celled stage) of the embryo (Fig. 7.128A, B).
Subsequent development of the upper two cells forms the root and
leaf, whereas the lower two cells give rise to the foot and shoot apex
(Fig. 7.128C, D).
With the development of the embryo the vegetative cells of the
surrounding gametophyte divides periclinally and form a two- to
three-layered sheath (calyptra) around the embryo. The primary root
grows vertically and establishes the sporeling in the soil. The young
sporophyte has a well-developed primary root and leaf.
The life cycle of Marsilea is shown in Fig. 7.129.

Morphological Nature of Sporocarp:

The morphological nature of the sporocarp of Marsilea has been
controversial. There are three views which can be grouped under the
following categories :
(i) Leaf segment theory or Laminar hypothesis,
(ii) Whole leaf theory or Petiolar hypothesis,
(iii) A phylogenetically synthesised new organ.
(i) Leaf Segment Theory or Laminar Hypothesis:
On the basis of the vascular supply to the peduncle as well as within
the sporocarp, this structure has been considered to be a modification
of a leaf segment rather than an entire leaf. This hypothesis was
supported from time to time by a number of scientists. Although the
scientists equated sporocarp with the lateral fertile part of a leaf, they
differed on the number of pinnae involved and their mode of
transformation into the sporocarp.
The Various Views:
According to Russow (1872), the sporocarp is made up of two leaflets
and their ventral surfaces containing sori are facing each other.
Bower (1889) interpreted sporocarp to be a modified unipinnately
compound leaf with rachis bearing two rows of pinnules which fused
laterally to form sporocarp (Fig. 7.130A, B).
According to Campbell (1892) the sporocarp is a folded pinnately
compound leaf where pinnae are fused to form the sporocarp.
Eames (1936) equated the sporocarp with the tip of the leaf whose
margins bend downwards (Fig. 7.130C-D). A wing-like shoulder
develops that protects the sori. The similarity between the course of
vascular traces in the sporocarp and the vasculature of the leaf is an
evidence in support of Eames’ hypothesis.
According to Smith (1955) the sporocarp is a modified pinna with two
abaxial sori. The infolding of the tvyo edges of the pinna with abaxial
sori along the midrib forms the sporocarp. In the axil of each
receptacle a flap-like indusium is formed (Fig. 7.130E).
Puri and Garg (1953) interpreted the sporocarp as a modified single
leaflet in which the number of pinnules corresponds to the number of
lateral bundles in the sporocarp (Fig. 7.130F).
(ii) Whole Leaf Theory or Petiolar Hypothesis:
Johnson (1933) is the proponent of this theory. According to this
hypothesis, the sporocarp is the modification of the whole leaf (lamina
and petiole) where the marginal cells of the leaf develop into sporangia
instead of leaflets. Evidence in support of this hypothesis is derived
from the development of secondary and even tertiary sporocarps on
peduncle of primary sporocarp.
(iii) A Phylogenetically Synthesized New Organ:
Bierhorst (1971) interpreted the sporocarp of Marsileaceae to be a
phylogenetically synthesized new organ which changed its
morphological nature in mid-ontogeny. According to Bierhorst the
sporocarp originates after the vegetative pinnae are formed. Each
sporocarp arises from a single initial with three cutting faces, a
condition unknown for any other fern pinna.
In the early stages of development, the marginal series of sporocarp
initials are comparable to the sporangial stalk initials of ferns. Thus,
there is a differential expression of genome where sporangial ontogeny
is expressed in early stage and pinna ontogeny in later stage.