Text

Systematic Position
Division: Lycopodiophyta
Class:

Isoetopsida

Order: Selaginellales
Family: Selaginellaceae
Genus: Selaginella
Habit and Habitat
Selaginella, with about 700 species, is cosmopolitan in
distribution (Banks, 2009). The species are commonly
known as spike moss or small club moss. Most of the
species inhabit damp and shaded forests of tropics,
but some (e.g., S. densa, S. rupestris, S. lepidophylla)
grow in xerophytic habitats, such as exposed rock
surfaces. S. oregano is an epiphyte that

Fig 1:

Habit of Selaginella growing in xerophytic

conditions

grows on tree trunks in tropical rain forests. Several

species of Selaginella are grown in gardens as
ornamentals (Rashid, 1999). Some xerophytic species
of Selaginella (e.g., S. lepidophylla, S. pilifefra) show
caespitose habit; they curl and become ball like during
dry season and again become green and fresh when
moisture is available. These are called resurrection
plants (Singh et al, 2000).

Fig 2: Selaginella showing caespitose habit

Selaginella

is

particularly

interesting

from

comparative evolutionary perspective because it has

retained

the

independent

but

water-dependent

gametophytic generation that is typical of all non-seed

plants. Because its gametophyte is not buried within
maternal tissues of the sporophyte, Selaginella is also
a useful experiment system for investigating how the
alternation of generations (the switch between haploid
gametophyte and diploid sporophyte) is regulated
(Banks, 2009)

The genus is represented in India by more than

70

species.

Among

these

species,

Selaginella

kraussiana, S. monospora, S. biformes, S. rupestris,

S.

megaphylla,

S.

bryopteris,

S.

ciliaris,

S.

chrysorhizos and S. pentagona are common.

Morphology
The

sporophytic

plant

body

of

Selaginella

is

differentiated into root, stem and leaves. Besides

some species also have rhizophores.
1.

Roots. The primary roots are ephemeral and the

adult plant has adventitious roots. The adventitious

roots usually have specific locations in relation to stem
dichotomies. In most of the creeping species with
dorsiventral stems (e.g., S. kraussiana, S. laevigata),
roots arise at or close to the point of dichotomy; in
species like S. rupestris and S. wallichii they arise at
the point of dichotomy as well as at other positions;
and in S. selaginoides and S. spinulosa they arise
from knot-like swellings present at the basal portion
of the stem.
The

roots

arise

endogenously

and

are

dichotomously branched. The dichotomies are at right

angles to each other. The main function of root is to
anchor the plant in the soil and absorb water and

mineral salts from the soil. Besides it form a passage

way for water and dissolved substances from the root
into the stem and also for foods from the stem down
into the root.

Fig 3: Adventitious roots in Selaginella

2.

Rhizophore. In some species of Selaginella,

many

long,

cylindrical,

unbranched

and

leafless

structures arise from the lower side of the stem at the

point of dichotomy. These grow vertically downward
and bear tuft of adventitious roots at their distal end.
They are known as rhizophores (Goebel 1905; Bower,
1935). The rhizophore may develop into a typical leafy
shoot under certain conditions.

Like a typical root it

grows downwards to the soil and absorbs water

through its tissues in a direction reverse of that in
which it has grown. It produces lateral endogenous

roots

and

helps

in

anchoring

the

plant

to

the

substratum.

Fig 4: Showing various organs like rhizophore,

cone and leaves in Selaginella
3.

Stem. The stem is erect and dichotomously

branched in the sub-genus Homoeophyllum, and

prostrate or sub-erect with lateral branching in the
sub-genus Heterophyllum.
The stem apex usually has a single well-defined apical
cell, but in S. oregano a group of meristematic cells
has been observed.
4.

Leaves.

The

leaves

of

Selaginella

are

microphyllus, sessile and simple. Their shape varies

from ovate to lanceolate. The leaf has a single
midvein

that

remains

unbranched

throughout

its

course. Most of the species have thin and soft leaves,

but

in

xerophytic

species

they

are

thick.

The

vegetative leaf as well as sporophyll, has a small

membranous projection on its adaxial (upper) surface,
close to the base. The projection is known as the
ligule. The basal part of the ligule has a distinct
hemispherical foot-like structure, called glossopodium.
It

is

composed

of

highly

vacuolated

thin-walled

tubular cells. The ligule is embedded at the base of

the leaf in a pit like structure, known as ligular pit.
The projected part of the ligule is only one cell in
thickness and is tongue-like (e.g., S. svogelli, S.
martensii). It develops precociously and matures long
before

its

associated

leaf.

Although

the

definite

function of ligule is not known, it has been suggested

that in some way they are associated with water
absorption

and

secretion,

and

thus

prevent

desiccation of the shoot. Some consider that the

ligules in Selaginella are concerned with upward
movement of inorganic solutes, and thus compensate
for smaller and less effective leaf primordia.
In the sub-genus Homoeophyllum, all leaves are
alike and spirally arranged. But the species belonging
to the sub-genus Heterophyllum, have two types of
leaves- two dorsal rows of small leaves (microphylls),
and two ventral rows of large leaves (megaphylls).

The leaves occur in pairs and the two leaves of a pair

are always unequal.
Anatomy
1.

Root.

A cross section of a root shows a simple structure.

The epidermis is made up of tangentially elongated
cells. In exposed roots, the outer wall of epidermal
cells is cutinized, but in roots that penetrate the
substratum, the epidermal cells are delicate and have
root hairs.
The cortex is usually homogenous, consisting of many
layers of thin parenchymatous cells. But in some
species the outer layers of the cortex become thickwalled and form hypodermis. In S. selaginoides, the
parenchymatous cells of the cortex show mycorrhizal
association. The innermost layer of the cortex forms
endodermis. In species like S. densa and S. rubella
endodermis is fairly distinct.
The central part of the root is occupied by a
protostele,

surrounded

by

1-3

layers

of

parenchymatous pericycle. The xylem, which forms

the central solid core of the stele, is monarch to
tetrarch and exarch. The phloem occurs in the form of
a ring around the xylem.
2.

Rhizophore.

The anatomy of the rhizophore resembles that of the

root. Some variations in the internal organization are
due to the fact that the rhizophore is an aerial
structure, whereas the root is a subterranean organ.
The epidermis is single layered and the outer wall of
the epidermal cells is covered with a thick layer of
cuticle. Root hairs, characteristic of roots, are absent
on rhizophores. The cortex is differentiated into an
outer sclerenchymatous and an inner relatively wide
parenchymatous zone. The innermost layer of the
cortex

forms

rhizophore

is

endodermis.
surrounded

The
by

protostele
a

of

the

parenchymatous

pericycle. Usually the stele is monarch and exarch, but

shows some variations. For example, in S. atro-viridis,
the

metaxylem

is

crescent-shaped

with

many

protoxylem strands on its concave side, and in S.

kraussiana the xylem is centrifugal.

Fig 5: Cross section of rhizophore

3. Stem.

Internally, the stem is more complex than the root.

The anatomy of the stem shows variations not only in
different species, but also within the same species
depending on stem diameter. A transverse section of
the

stem

shows

epidermis,

cortex

and

central

cylinder.

Fig 6: Internal structure of Selaginella stem

The epidermis is the outermost unistratose layer. The
outer walls of the epidermal cells are highly cutinized.
The epidermis is devoid of stomata and hairs.
The cortex is usually composed of compactly arranged
parenchymatous cells without intercellular spaces. But
in mature stems of many species outer layers of
cortex become partially sclerenchymatous, forming a
tough hypodermis. In xerophytic species (e.g., S.

rupestris, S. lepidophylla), most part of the cortex is

composed of thick-walled cells.
A

distinctive

feature

of

Selaginella

stem

is

the

presence of radially elongated endodermal cells, called

trabeculae. They have characteristic casparian bands
on

their

lateral

walls.

Due

to

the

presence

of

trabeculae, the central stele is separated from the

cortex by large air spaces. In fact, in a transverse
section the stele appears suspended in an axial air
column with the help of trabeculae. The air spaces
develop due to more rapid expansion of the cortical
region than the stele. This differential growth also
results in radial stretching of some endodermal cells.
Xerophytic species of Selaginella, however, do not
have trabeculae.

Fig 7: Showing trabeculae in Selaginella stem

The number of steles in the stem shows considerable
variation

in

different

species

of

Selaginella.

For

example, the stem is monostelic in S. spinulosa and S.

flabellata, distelic in S. kraussiana, and polystelic

(with 12-16 steles) in S. laevigata. Besides, the
number of steles may also vary within different parts
of

the

same

plant.

For

example,

the

creeping

branches of S. braunii are distelic, whereas the erect

branches are monostelic; and in S. lyalli, the creeping
branches are distelic and the erect branches are
polystelic.
The stele is surrounded by a single-layered pericycle.
The shape and structure of the stele is also variable. It
is circular in S. kraussiana and flat or ribbon-like in S.
viridangula and S. vagelii. Most of the species have a
protostele with a solid xylem core surrounded by
phloem, but S. laevigata var. lyalii has a siphonostele.
The xylem is usually monarch (S. kraussiana), or
diarch (S. selaginoides). It usually consists of only
tracheids; the protoxylem tracheids have annular or
helical thickenings; whereas the metaxylem tracheids
show scalariform thickenings. In S. oregana, S. densa
and S. rupestris, however, the xylem has true vessels
with

transverse

perforation

plates.

Although

secondary growth is absent, some secondary xylem

elements have been found in the basal part of the
stem of S. selaginoides.
4

Leaf.

Both, the upper and the lower epidermis of the leaf

are unistratose. The epidermal cells have chloroplasts.
The leaves are mostly amphistomatic, but sometimes
they are hypostomatic, as in S. martensii. Stomata
are distributed mostly in the midrib region.
The mesophyll consists of loosely arranged thinwalled cells, with many small or large intercellular
spaces.

It

is

usually

made

up

of

only

spongy

parenchyma, but occasionally a distinct palisade layer

may be present towards the morphological upper side.
A mesophyll cell has 1-8 cup shaped chloroplasts,
which have many spindle shaped pyrenoid-like bodies.
The leaf has a median vascular bundle surrounded
by

distinct

bundle

sheath.

The

xylem,

which

occupies the central part of the bundle, consists of

only tracheids with annular or spiral thickenings. It is
surrounded by phloem.
Reproduction
The

sporophyte

of

Selaginella

reproduces

vegetatively and by spores.

Vegetative reproduction
Vegetative propagation in Selaginella takes place
by

tubers,

fragmentation.

bulbils,

dormant

buds

and

by

In S. rupestris, prostrate branches produce roots

during

favorable

conditions.

These

root

bearing

prostrate branches separate from the parent plant and

grow into new sporophytes.
Species like S. chrysorhizos and S. chrysocaulos
propagate with the help of tubers and bulbils. The
tubers may be aerial, developing at the apices of
aerial

branches

(e.g.,

S.

chrysocaulos)

or

subterranean (e.g., S. chrysorhizos). During favorable

conditions

the

tuber

germinates

into

new

sporophyte. Aerial branches of S. chrysocaulos also

bear some dormant (resting) buds which grow into
new plants during favorable conditions.

Fig 8: Tubers in Selaginella

Reproduction by spores
Selaginella

is

heterosporous

pteridophyte;

it

produces two types of spores megaspores and

microspores.

The

megaspores

form

female

gametophytes on germination and the microspores

give rise to male gametophytes. The sporangia are

strictly dimorphic, i.e., micro and megaspores are
formed in separate sporangia. The sporangia bearing
microspores are called microsporangia, and those
bearing megaspores as megasporangia. There are
many microspores in a microsporangium, while each
megasporangium usually has 1-4 (or rarely more)
megaspores. The megaspores are much larger than
the microspores.
The sporangia are borne singly in the axils of
sporophylls. The sporophyll-bearing micro-sporangium
is

called

microsporophyll,

and

the

one

with

megasporangium is known as megasporophyll. The

sporophylls are spirally arranged around a central axis
to form a strobilus.

Fig 9: Parts of strobilus showing megaspores and

microspores in Selaginella

Strobilus or cone. In most of the species of

Selaginella, sporophylls are aggregated at the apex of
the main stem and its branches in definite loose or
compact cones, called strobili (singular = strobilus).
The size of the strobilus varies from 5mm to 6-7 cm.
It is often inconspicuous due to its small size, and
similarity between sporophylls and vegetative leaves.
Usually a branch terminates in strobilus, but in species
like S. cuspidata and S. patula, vegetative growth of
the branch may continue beyond the strobilus. In S.
erythropus, a second strobilus is produced on the
fertile branch after an intervening vegetative region.
Thus, in this species sporophylls and vegetative leaves
occur in alternate segments.
Distribution of micro and megasporangia in
strobilus. In most of the species of Selaginella, both
micro and megasporangia are found within the same
strobilus. Their distribution is specific. For example, in
S.

selaginoides,

S.

rupestris

and

S.

helvetica,

megasporangia are present in the basal part and

microsporangia in the upper part of the strobilus; in S.
kraussiana there is only a single megasporangium at
the

base

of

the

strobilus,

and

the

rest

are

microsporangia; and in S. inaequalifolia one side of

the strobilus bears only megasporangia, and the other

microsporangia. In S. martensii and S. caulescens,

mega-and microsporangia do not show any definite
arrangement. In S. selaginoides, a series of basal
sporangia are non- functional. The two types of spores
are never present within the same sporangium.
In S. gracilis and S. atroviridis, strobili are
monosporangiate, i.e. micro and megasporangia
are borne in separate strobili.
Development of sporangium. The initial stages of
the development of micro and megasporangium are
similar. Both develop from the transverse row of initial
cells, i.e. the development is of eusporangiate type.
The sporangial initials divide periclinally, establishing
outer jacket initials and inner archesporial initials. The
archesporial initials undergo repeated anticlinal and
periclinal divisions forming a mass of sporogenous
cells. Simultaneous divisions also occur in the jacket
initials and the derivatives eventually form a twolayered sporangial jacket. The cells of the outermost
layer of the sporogenous tissue (adjacent to the inner
wall layer) form a nutritive layer, known as tapetum.
The tapetal layer disintegrates as spores mature.
The last generation of sporongenous cells functions
as

spore

mother

cells.

The

micro

and

megasporangium differ in subsequent development.

Further development of microsporangium. In

microsporangium about 80-90% spore mother cells
are functional, and behave as microspore mother
cells. The remaining spore mother cells degenerate
and form a viscous nourishing fluid. The functional
spore mother cells undergo meiosis and form haploid
microspores,

which

are

arranged

in

tetrahedral

tetrads.
Further development of megasporangium. In
megasporangium, all spore mother cells but one,
degenerate. The functional spore mother cell behaves
as megaspore mother cell. It divides meiotically
forming

four

tetrahedrally

arranged

haploid

megaspores. All the four megaspores derived from a

megaspore mother cell may not always be functional.
For example, in S. sulcata only one, and in S.
rupestris two megaspores are functional. Sometimes
there are more than one megaspore mother cells in a
megasporangium

and

in

such

cases

the

megasporangium has 8 or more megaspores. The

megaspores are much larger than microspores.
The expression of maleness or femaleness is not
genetically determined; it appears to be influenced by
the nutritional factor, and the specific environment in
which the sporangium develops.

Mature sporangium.

Mature sporangia are

stalked structures, with a two-layered sporangial

jacket. The cells of outer jacket layer are elongated
and

contain

chloroplasts.

The

micro

and

mega

sporangia differ in shape, size and colour. The

microsporangia are slightly elongated, yellow, red or
orange in colour. The megasporangia are larger and
paler and assume the shape dictated by the enlarging
megaspores within.
The mature sporangium dehisces along the line of
dehiscence present at its distal end and oriented
transverse to the axis of the sporophyll. Structural
modification of the surface cells along this line and at
its flanks results in splitting of the distal part of the
sporangium into two valves. The lower cup-shaped
part of the sporangium shrinks on drying and throws
out spores violently.
Gametophyte
The spore is the mother cell of the gametophytic
generation.

As

mentioned

earlier,

Selaginella

is

heterosporous and produces two types of spores- the

smaller microspore and the larger megaspores. This
difference in the size of the spores is related to their
fate and function; microspores develop into male

gametophyte

and

megaspores

into

female

gametophytes.
In Selaginella both microspores and megaspores
begin to germinate while still inside the sporangium
(i.e., they germinate in situ). Thus, spores are shed at
multicellular stage.
Microspores

and

development

of

male

are

small,

gametophyte
Microspores:
spherical

The

structures,

microspores

ranging

0.015-0.06

mm

in

diameter. A microspore is surrounded by a thick

ornamented exine and a relatively thin intine. The
ornamentations

in

the

exine

may

be

papillate,

echinulate or granulate. The spore has a single

haploid nucleus and granular cytoplasm, rich in fatty
substances. The fats probably provide food to the
developing male gametophytes as spores contain no
chlorophyll.
Development

of

male

gametophyte: The

microspores germinate inside the microsporangium

and are shed at 13- celled stage. The first division of
the microspore is asymmetrical and as a result a small
lenticular prothallial cell and a large antheridial initial
is established. The prothallial cell does not divide

further and the entire sporangium develops from the

antheridial initial.
The first division of the antheridial initial is nearly
at right angles to the prothallial cell. It results in the
formation of two antheridial cells of almost equal size.
Both these cells divide by a vertical wall to produce a
group

of

four

cells.

Thus,

at

this

stage

the

gametophyte consists of five cells (four cells derived

from the antheridial initial and a prothallial cell). The
two basal cells, derived from the antheridial initial, do
not divide further, whereas the upper two daughter
cells divide repeatedly and form ten cells. At this
stage the gametophyte has 13 cells (10 cells derived
from the upper daughter cells of the antheridial
initial,2 basal daughter cells and 1 prothallial cell).Of
these,

four

central

cells

function

as

primary

androgonial cells and eight peripheral ones as jacket

cells.

The

male

gametophyte

is

shed

from

the

microsporangium at 13-celled stage. It enters into

partially

opened

megasporangium

where

further

development of the male gametophyte takes place in

close

proximity

of

the

developing

female

gametophyte. In some species, it is believed, that

further developed of the male gametophyte takes
place in the soil.

The four central primary androgonial cells of the

male gametophyte divide repeatedly forming a mass
of 128-256 antherozoid mother cells or androcytes.
Each androcyte metamorphoses into a spindle-shaped
biflagellate

antherozoid.

The

antherozoids

of

Selaginella are perhaps the smallest amongst the

vascular plant.
With the formation of antherozoids, the jacket
cells decompose and form a mucilaginous substance.
The antherozoids float in this substance. Until this
stage the male gametophyte is completely enclosed
within the wall of the microspore. Thus it is entirely
endosporic and extremely reduced structure. Unlike
other

pteridophytes,

vegetative

prothalli

are

not

formed in Selaginella. The gametophyte is not set free

and is dependent on the parent sporophyte for
nutrition.
Megaspore

and

development

of

female

gametophyte
Megaspores: Megaspores are much larger than
the microspores. Their diameter varies from 0.15 to
0.5 mm. Usually all megaspores in a megasporangium
are approximately of the same size, but in S.
molliceps one megaspore is larger than the other
three, and in S. stenophylla there are two large and

two small megaspores. The megaspores are also

arranged in tetrahedral tetrads. The wall of the
megaspore is differentiated into an outer massive
exine and an inner thin intine, but in S. rupestris and
S. apus it is differentiated into three distinct layersthe outer exospore, the middle mesospore and the
inner endospore. The megaspore has a single haploid
nucleus, surrounded by granular cytoplasm, rich in
fatty substance.
Development of female gametophyte: Like
male gametophyte, the development of the female
gametophyte of Selaginella also begins while it is still
within the megasporangium. In S. kraussiana, the
gametophyte is liberated from the megasporangium
after the first archegonium is differentiated, whereas
in S. rupestris and S. apus it is retained in the
megasporangium

even

after

the

development

of

embryo has started. However, in S. spinulosa and S.

helvitica the development of female gametophyte
starts only after the megaspore is shed from the
sporangium.
Immediately

after

the

development

of

female

gametophyte initiates, a large vacuole appears in the

centre of the megaspore and as a result the cytoplasm
is pushed along the spore wall in the form of thin

membrane. There is considerable enlargement of the

megaspore. The outer spore wall (exospores) grows
more rapidly than the mesospore and endospore,
consequently a large gap is formed in between the
exospore and mesospore. At this stage, the exospore
is attached to the mesospore only at one point. The
space between the exospore and the mesospore is
filled with a homogenous liquid.
The

haploid

nucleus

of

the

megaspore

divides

repeatedly without any wall formation. The free nuclei

are unequally distributed in the peripheral cytoplasm;
they are clustered beneath the triradiate ridge of the
spore and sparsely distributed elsewhere. Now, wall
formation begins in the apical region and a lensshaped pad of small cells is formed at the apical end.
It

is

separated

gametophyte

by

from
a

the

rest

distinct

of

the

female

diaphragm.

The

cytoplasmic layer becomes thicker gradually and

pushes the mesospore outward. As a result the
mesospore again comes in contact with the exospores.
With the increase in the amount of cytoplasm, the
central vacuole diminishes and eventually disappears.
The part of the gametophyte below the diaphragm is
multinucleate

in

early

stages

but

becomes

multicellular as wall formation proceeds inward. At

this stage, the spore wall ruptures along the triradiate

ridge exposing the apical cellular pad. The exposed
part

of

the

female

gametophyte

may

develop

chloroplasts but the photosynthetic ability of this part

is of limited importance as food for the developing
embryo is stored in the lower multicellular part of the
gametophyte.

Many

rhizoids

develop

from

the

exposed part of the gametophyte. They attach the

gametophyte to the substratum and also help in
absorption of water.
Development

of

archegonia:

Archegonia

develop from the apical tissue of the gametophyte. All

superficial cells of this tissue have the potential of
forming archegonia. The archegonial initial divides
periclinally into a primary cover cell and a central cell.
The primary cover cell divides by two vertical divisions
at right angles to each other and forms four neck
initials. The neck initials divide transversely so as to
form eight neck cells, arranged in two tiers of four
each, in the meantime, the central cell divides by a
periclinal wall and an outer primary neck cell and an
inner primary venter cell is established. The former
does not divide further and directly functions as neck
canal cell, whereas the latter divides transversely into
a venter canal cell and an egg.

The mature archegonium of Selaginella has two cell

long neck (consisting of eight cells in two tiers of four
each), a neck canal cell, a venter canal cell and an
egg. The four terminal cells of the neck project
beyond the surface of the gametophyte as asymmetric
nipples. Rest of the archegonium remains embedded
in the tissue of the gametophyte

Fig 10: Spores and their fate

Fertilization
Fertilization usually takes place after the
megasporangium has fallen on the soil, but in some
species it may occur while the female gametophyte is
still within the sporangium. Just before fertilization,
the neck cells of the archegonium separate from each
other

and

form

passage

for

the

entry

of

antherozoids.

After

liberation

from

the

male

gametophyte, antherozoids swim in rain or dew water

and

reach

the

archegonia.

Usually

only

one

antherozoid enters into an archegonium and fuses

with the egg to form a diploid zygote.
Some species of Selaginella (e.g., S. rupestris, S.
apoda)

show

seed

habit. In

these

species, the

sporangium has only a single megaspore and at

maturity of the archegonium the spore wall ruptures,
but the developing female gametophyte does not
come out of the spore wall. The developing male
gametophyte, when shed from the microsporangium
(present in the distal part of the strobilus) lands on
the partially open megasporangium. Thus, at this
stage, both the male and the female gametophytes lie
within the megasporangium. As such fertilization and
embryo

development

take

place

inside

the

megasporangium. The sporangium is shed after the

development of root and primary shoot of the new
sporophyte. This feature is of considerable importance
from the point of view of seed habit because when the
megaspore with young sporophyte is shed, it has all
typical characters of a seed.
Development of embryo

The

diploid

sporophytic

zygote

is

generation.

the

mother

cell

of

the

It

divides

transversely,

establishing an epibasal (upper) suspensor cell and a

hypobasal (lower) embryonic cell.

As development

proceeds, the suspensor cell repeatedly divides to

form a suspensor, which pushes the developing
embryo deep into the female gametophyte. The rest
of the embryo develops from the embryonic cell. It
divides by two vertical walls at right angles to each
other, and thus a four-celled embryo is formed. One
of the four cells of the embryo divides by an oblique
vertical wall, and thus an apical cell with three cutting
faces is established. This eventually functions as the
apical cell of the embryonic shoot.The remaining three
cells of the 4-celled embryo and the sister cell of the
apical cell (i.e., total four cells) divide transversely to
form two tiers of four cells each. The cells of both the
tiers divide irregularly forming a multicellular embryo.
Usually the cells of lower tier divide more rapidly than
the upper tier and due to this differential growth the
entire embryo apex rotates at 1800 and emerges
through the apical part of the gametophyte. The
derivatives of the lower tier form the foot. At first the
foot grows on one side but eventually comes to lie
opposite the suspensor. The foot acts as a haustorial

organ; its main function is to absorb nutrition for the

developing sporophyte from the female gametophyte.
At this stage, a superficial cell in each of the two
diagonally

opposed

quadrants

of

the

upper

tier

differentiates as the apical cell of a foliar appendage,

which eventually forms a cotyledon. In the axil of each
cotyledon a ligule develops.
The part of the embryo immediately posterior to
cotyledons develops into hypocotyledonary

part of

the stem. The stem grows with the help of the apical
cell of the embryo. After the formation of cotyledons
and stem, the apical cell of the root differentiates on
the lateral surface of the foot. The derivatives of this
cell

develop

into

root-like

structure,

called

rhizophore. Roots, in fact, develop at the apex of the

rhizophore. In early stages of development the young
sporophyte is attached to the megaspore and derives
its food from the female gametophyte with the help of
its foot. But after the establishment of root and stem,
the sporophyte becomes independent.

Fig 11: General life-cycle of Selaginella

Medicinal uses
Many species of Selaginella have been used as
traditional

medicines.

In

India,

S.

bryopteris

is

referred to as Sanjeevanione that infuses lifefor its

medicinal properties (Sah et al. 2005). In Columbia,
S. articulata is used to treat snakebites and neutralize
Bothrops atrox venom. Throughout southern China,
Selaginella is used as a popular herb for the treatment
of various ailments (Lin and Kan, 1990; Pan et al
2001 and Maa et al 2003). Although most reports of
the medicinal uses of Selaginella are anecdotal,
researchers have begun to identify and characterize
the active compounds in Selaginella extracts (Kang et

al. 2004; Chen et al. 2005 and Yin et al. 2005).

Among the best characterized are uncinoside A and
uncinoside B, biflavonoids that have potent antiviral
activities against respiratory syncytial virus (Ma et al
2003). Other biflavonoids from S. tamariscina inhibit
the induction of nitric oxide (NO) and prostaglandins
(Pokharel et al 2006; Woo et al 2006; Yang et al,
2006), which are involved in the pathogenesis of
some cancers (Lala and Chakraborty, 2001; Zha et al
2004). The biflavone ginkgetin from S. moellendorffii
selectively inhibits the growth of some cancer cells by
inducing apoptosis (Sun et al 1997; Su et al 2000).