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Systematic Position
Division: Lycopodiophyta


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


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





comparative evolutionary perspective because it has





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






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







chrysorhizos and S. pentagona are common.







differentiated into root, stem and leaves. Besides
some species also have rhizophores.

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.






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

Rhizophore.mineral salts from the soil. Fig 3: Adventitious roots in Selaginella 2. Bower. These grow vertically downward and bear tuft of adventitious roots at their distal end. cylindrical. It produces lateral endogenous . In some species of Selaginella. 1935). They are known as rhizophores (Goebel 1905. The rhizophore may develop into a typical leafy shoot under certain conditions. many long. unbranched and leafless structures arise from the lower side of the stem at the point of dichotomy. 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. 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.

roots and helps in anchoring the plant to the substratum. 4. The leaves of Selaginella are microphyllus. The leaf has a single midvein that remains unbranched throughout its course. cone and leaves in Selaginella 3. but in xerophytic species they are thick. and prostrate or sub-erect with lateral branching in the sub-genus Heterophyllum. but in S. Fig 4: Showing various organs like rhizophore. Most of the species have thin and soft leaves. Stem. The . sessile and simple. The stem apex usually has a single well-defined apical cell. Their shape varies from ovate to lanceolate. The stem is erect and dichotomously branched in the sub-genus Homoeophyllum. oregano a group of meristematic cells has been observed. Leaves.

The basal part of the ligule has a distinct hemispherical foot-like structure.two dorsal rows of small leaves (microphylls). S. But the species belonging to the sub-genus Heterophyllum. Although the definite function of ligule is not known. Some consider that the ligules in Selaginella are concerned with upward movement of inorganic solutes. The projection is known as the ligule. close to the base.g.vegetative leaf as well as sporophyll. S. The projected part of the ligule is only one cell in thickness and is tongue-like (e. In the sub-genus Homoeophyllum. it has been suggested that in some way they are associated with water absorption and secretion. martensii). has a small membranous projection on its adaxial (upper) surface. all leaves are alike and spirally arranged. The ligule is embedded at the base of the leaf in a pit like structure. and two ventral rows of large leaves (megaphylls).. It is composed of highly vacuolated thin-walled tubular cells. It develops precociously and matures long before its associated leaf. and thus compensate for smaller and less effective leaf primordia. . known as ligular pit. svogelli. and thus prevent desiccation of the shoot. have two types of leaves. called glossopodium.

The xylem. the epidermal cells are delicate and have root hairs. A cross section of a root shows a simple structure. surrounded by 1-3 layers of parenchymatous pericycle. The innermost layer of the cortex forms endodermis. selaginoides. The phloem occurs in the form of a ring around the xylem. the parenchymatous cells of the cortex show mycorrhizal association. Anatomy 1. is monarch to tetrarch and exarch. . The epidermis is made up of tangentially elongated cells. Root. Rhizophore.The leaves occur in pairs and the two leaves of a pair are always unequal. consisting of many layers of thin parenchymatous cells. densa and S. which forms the central solid core of the stele. In S. 2. the outer wall of epidermal cells is cutinized. but in roots that penetrate the substratum. But in some species the outer layers of the cortex become thickwalled and form hypodermis. The central part of the root is occupied by a protostele. In species like S. The cortex is usually homogenous. In exposed roots. rubella endodermis is fairly distinct.

The cortex is differentiated into an outer sclerenchymatous and an inner relatively wide parenchymatous zone. For example. surrounded The by protostele a of the parenchymatous pericycle. kraussiana the xylem is centrifugal. The epidermis is single layered and the outer wall of the epidermal cells is covered with a thick layer of cuticle.The anatomy of the rhizophore resembles that of the root. Fig 5: Cross section of rhizophore 3. atro-viridis. Root hairs. The innermost layer of the cortex forms rhizophore is endodermis. but shows some variations. Stem. Some variations in the internal organization are due to the fact that the rhizophore is an aerial structure. the metaxylem is crescent-shaped with many protoxylem strands on its concave side. whereas the root is a subterranean organ. in S. Usually the stele is monarch and exarch. characteristic of roots. are absent on rhizophores. and in S. .

cortex and central cylinder. In xerophytic species (e.g. The epidermis is devoid of stomata and hairs. forming a tough hypodermis. The cortex is usually composed of compactly arranged parenchymatous cells without intercellular spaces. A transverse section of the stem shows epidermis. Fig 6: Internal structure of Selaginella stem The epidermis is the outermost unistratose layer. The anatomy of the stem shows variations not only in different species. the stem is more complex than the root. But in mature stems of many species outer layers of cortex become partially sclerenchymatous.Internally. . S. The outer walls of the epidermal cells are highly cutinized. but also within the same species depending on stem diameter..

For example. most part of the cortex is composed of thick-walled cells. Xerophytic species of Selaginella. A distinctive feature of Selaginella stem is the presence of radially elongated endodermal cells. however. In fact. This differential growth also results in radial stretching of some endodermal cells. in a transverse section the stele appears suspended in an axial air column with the help of trabeculae. Fig 7: Showing trabeculae in Selaginella stem The number of steles in the stem shows considerable variation in different species of Selaginella. . lepidophylla). S. The air spaces develop due to more rapid expansion of the cortical region than the stele. They have characteristic casparian bands on their lateral walls. the central stele is separated from the cortex by large air spaces. do not have trabeculae. called trabeculae.rupestris. Due to the presence of trabeculae. the stem is monostelic in S. spinulosa and S.

some secondary xylem elements have been found in the basal part of the stem of S. 4 Leaf. rupestris. the number of steles may also vary within different parts of the same plant. kraussiana and flat or ribbon-like in S. lyalii has a siphonostele. whereas the metaxylem tracheids show scalariform thickenings. It usually consists of only tracheids. The xylem is usually monarch (S. S. laevigata var. the creeping branches of S. The stele is surrounded by a single-layered pericycle. For example. Although secondary growth is absent. the protoxylem tracheids have annular or helical thickenings. the xylem has true vessels with transverse perforation plates. selaginoides. but S. lyalli. selaginoides). braunii are distelic. . and in S. and polystelic (with 12-16 steles) in S. whereas the erect branches are monostelic. The shape and structure of the stele is also variable. kraussiana). vagelii. Besides. or diarch (S.flabellata. In S. viridangula and S. laevigata. distelic in S. It is circular in S. oregana. kraussiana. the creeping branches are distelic and the erect branches are polystelic. densa and S. however. Most of the species have a protostele with a solid xylem core surrounded by phloem.

dormant buds and by . consists of only tracheids with annular or spiral thickenings. as in S. with many small or large intercellular spaces. A mesophyll cell has 1-8 cup shaped chloroplasts.Both. which occupies the central part of the bundle. It is surrounded by phloem. the upper and the lower epidermis of the leaf are unistratose. bulbils. The xylem. fragmentation. martensii. Vegetative reproduction Vegetative propagation in Selaginella takes place by tubers. Stomata are distributed mostly in the midrib region. The leaf has a median vascular bundle surrounded by a distinct bundle sheath. It is usually made up of only spongy parenchyma. The epidermal cells have chloroplasts. Reproduction The sporophyte of Selaginella reproduces vegetatively and by spores. but occasionally a distinct palisade layer may be present towards the morphological upper side. which have many spindle shaped pyrenoid-like bodies. The leaves are mostly amphistomatic. The mesophyll consists of loosely arranged thinwalled cells. but sometimes they are hypostomatic.

prostrate branches produce roots during favorable conditions. The megaspores form female gametophytes on germination and the microspores .g. chrysocaulos propagate with the help of tubers and bulbils. rupestris. chrysorhizos and S. S.In S.g. chrysocaulos) or subterranean (e. developing at the apices of aerial branches (e. The tubers may be aerial.. These root bearing prostrate branches separate from the parent plant and grow into new sporophytes. it produces two types of spores –megaspores and microspores. S. Fig 8: Tubers in Selaginella Reproduction by spores Selaginella is a heterosporous pteridophyte. Species like S. Aerial branches of S. During favorable conditions the tuber germinates into a new sporophyte.. chrysocaulos also bear some dormant (resting) buds which grow into new plants during favorable conditions. chrysorhizos).

e. The sporangia bearing microspores are called microsporangia. while each megasporangium usually has 1-4 (or rarely more) megaspores.. and those bearing megaspores as megasporangia. There are many microspores in a microsporangium. The sporangia are strictly dimorphic. micro and megaspores are formed in separate sporangia. The sporangia are borne singly in the axils of sporophylls. The sporophyll-bearing micro-sporangium is called microsporophyll. i. Fig 9: Parts of strobilus showing megaspores and microspores in Selaginella . The sporophylls are spirally arranged around a central axis to form a strobilus. The megaspores are much larger than the microspores.give rise to male gametophytes. and the one with megasporangium is known as megasporophyll.

In S. called strobili (singular = strobilus). inaequalifolia one side of the strobilus bears only megasporangia. rupestris and S. and the other . For example. sporophylls are aggregated at the apex of the main stem and its branches in definite loose or compact cones. Their distribution is specific. The size of the strobilus varies from 5mm to 6-7 cm. helvetica. and similarity between sporophylls and vegetative leaves. and the rest are microsporangia. In most of the species of Selaginella. Distribution of micro and megasporangia in strobilus. It is often inconspicuous due to its small size. selaginoides. vegetative growth of the branch may continue beyond the strobilus. in this species sporophylls and vegetative leaves occur in alternate segments. but in species like S.Strobilus or cone. in S. cuspidata and S. S. In most of the species of Selaginella. a second strobilus is produced on the fertile branch after an intervening vegetative region. kraussiana there is only a single megasporangium at the base of the strobilus. and in S. megasporangia are present in the basal part and microsporangia in the upper part of the strobilus. Usually a branch terminates in strobilus. both micro and megasporangia are found within the same strobilus. erythropus. patula. in S. Thus.

i. The two types of spores are never present within the same sporangium.e. caulescens. In S. martensii and S. the development is of eusporangiate type. i. The initial stages of the development of micro and megasporangium are similar. mega-and microsporangia do not show any definite arrangement. selaginoides.e. 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. The micro and megasporangium differ in subsequent development.functional. strobili are monosporangiate. Development of sporangium. atroviridis. a series of basal sporangia are non. known as tapetum. gracilis and S. establishing outer jacket initials and inner archesporial initials. Both develop from the transverse row of initial cells. The tapetal layer disintegrates as spores mature. In S. The archesporial initials undergo repeated anticlinal and periclinal divisions forming a mass of sporogenous cells.microsporangia. . The sporangial initials divide periclinally. In S. micro and megasporangia are borne in separate strobili. The last generation of sporongenous cells functions as spore mother cells.

The megaspores are much larger than microspores. All the four megaspores derived from a megaspore mother cell may not always be functional. . all spore mother cells but one. and the specific environment in which the sporangium develops. In microsporangium about 80-90% spore mother cells 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.Further development of microsporangium. and behave as microspore mother cells. The expression of maleness or femaleness is not genetically determined. rupestris two megaspores are functional. it appears to be influenced by the nutritional factor. degenerate. which are arranged in tetrahedral tetrads. The remaining spore mother cells degenerate and form a viscous nourishing fluid. The functional spore mother cells undergo meiosis and form haploid microspores. It divides meiotically forming four tetrahedrally arranged haploid megaspores. Further development of megasporangium. In megasporangium. The functional spore mother cell behaves as megaspore mother cell. For example. in S. and in S. sulcata only one.

the smaller microspore and the larger megaspores. The cells of outer jacket layer are elongated and contain chloroplasts. size and colour. The mature sporangium dehisces along the line of dehiscence present at its distal end and oriented transverse to the axis of the sporophyll. The megasporangia are larger and paler and assume the shape dictated by the enlarging megaspores within. Mature sporangia are stalked structures. microspores develop into male . Gametophyte The spore is the mother cell of the gametophytic generation. 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. red or orange in colour. The microsporangia are slightly elongated. Selaginella is heterosporous and produces two types of spores. As mentioned earlier. The lower cup-shaped part of the sporangium shrinks on drying and throws out spores violently. This difference in the size of the spores is related to their fate and function.Mature sporangium. The micro and mega sporangia differ in shape. with a two-layered sporangial jacket. yellow.

The ornamentations in the exine may be papillate. rich in fatty substances.gametophyte and megaspores into female gametophytes. gametophyte Microspores: spherical The structures. A microspore is surrounded by a thick ornamented exine and a relatively thin intine.015-0.. The first division of the microspore is asymmetrical and as a result a small lenticular prothallial cell and a large antheridial initial is established. echinulate or granulate. The prothallial cell does not divide . The fats probably provide food to the developing male gametophytes as spores contain no chlorophyll.celled stage. microspores ranging 0. Microspores and development of male are small.e. they germinate in situ). Development of male gametophyte: The microspores germinate inside the microsporangium and are shed at 13. In Selaginella both microspores and megaspores begin to germinate while still inside the sporangium (i. The spore has a single haploid nucleus and granular cytoplasm. Thus.06 mm in diameter. spores are shed at multicellular stage.

At this stage the gametophyte has 13 cells (10 cells derived from the upper daughter cells of the antheridial initial. at this stage the gametophyte consists of five cells (four cells derived from the antheridial initial and a prothallial cell). it is believed. It enters into partially opened megasporangium where further development of the male gametophyte takes place in close proximity of the developing female gametophyte. 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.further and the entire sporangium develops from the antheridial initial. The two basal cells.2 basal daughter cells and 1 prothallial cell). that further developed of the male gametophyte takes place in the soil. derived from the antheridial initial. . four central cells function as primary androgonial cells and eight peripheral ones as jacket cells. do not divide further. The male gametophyte is shed from the microsporangium at 13-celled stage. In some species. whereas the upper two daughter cells divide repeatedly and form ten cells.Of these. The first division of the antheridial initial is nearly at right angles to the prothallial cell.

Thus it is entirely endosporic and extremely reduced structure. Each androcyte metamorphoses into a spindle-shaped biflagellate antherozoid. but in S. The antherozoids float in this substance. With the formation of antherozoids.15 to 0.The four central primary androgonial cells of the male gametophyte divide repeatedly forming a mass of 128-256 antherozoid mother cells or androcytes. The antherozoids of Selaginella are perhaps the smallest amongst the vascular plant. and in S.5 mm. molliceps one megaspore is larger than the other three. the jacket cells decompose and form a mucilaginous substance. Their diameter varies from 0. Usually all megaspores in a megasporangium are approximately of the same size. Megaspore and development of female gametophyte Megaspores: Megaspores are much larger than the microspores. The gametophyte is not set free and is dependent on the parent sporophyte for nutrition. Until this stage the male gametophyte is completely enclosed within the wall of the microspore. stenophylla there are two large and . vegetative prothalli are not formed in Selaginella. Unlike other pteridophytes.

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

wall formation begins in the apical region and a lensshaped pad of small cells is formed at the apical end. the central vacuole diminishes and eventually disappears. At this stage. With the increase in the amount of cytoplasm. The outer spore wall (exospores) grows more rapidly than the mesospore and endospore. At . they are clustered beneath the triradiate ridge of the spore and sparsely distributed elsewhere. There is considerable enlargement of the megaspore. The part of the gametophyte below the diaphragm is multinucleate in early stages but becomes multicellular as wall formation proceeds inward. The cytoplasmic layer becomes thicker gradually and pushes the mesospore outward. the exospore is attached to the mesospore only at one point.membrane. The space between the exospore and the mesospore is filled with a homogenous liquid. It is separated gametophyte by from a the rest distinct of the female diaphragm. consequently a large gap is formed in between the exospore and mesospore. Now. As a result the mesospore again comes in contact with the exospores. The haploid nucleus of the megaspore divides repeatedly without any wall formation. The free nuclei are unequally distributed in the peripheral cytoplasm.

this stage. arranged in two tiers of four each. They attach the gametophyte to the substratum and also help in absorption of water. in the meantime. Many rhizoids develop from the exposed part of the gametophyte. All superficial cells of this tissue have the potential of forming archegonia. Development of archegonia: Archegonia develop from the apical tissue of the gametophyte. 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. The primary cover cell divides by two vertical divisions at right angles to each other and forms four neck initials. The archegonial initial divides periclinally into a primary cover cell and a central cell. the central cell divides by a periclinal wall and an outer primary neck cell and an inner primary venter cell is established. whereas the latter divides transversely into a venter canal cell and an egg. the spore wall ruptures along the triradiate ridge exposing the apical cellular pad. . The neck initials divide transversely so as to form eight neck cells. The former does not divide further and directly functions as neck canal cell.

The mature archegonium of Selaginella has two cell long neck (consisting of eight cells in two tiers of four each). a neck canal cell. 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. a venter canal cell and an egg. The four terminal cells of the neck project beyond the surface of the gametophyte as asymmetric nipples. the neck cells of the archegonium separate from each other and form a passage for the entry of . but in some species it may occur while the female gametophyte is still within the sporangium. Just before fertilization.

S. Usually only one antherozoid enters into an archegonium and fuses with the egg to form a diploid zygote. This feature is of considerable importance from the point of view of seed habit because when the megaspore with young sporophyte is shed. Thus.g.antherozoids. The developing male gametophyte. As such fertilization and embryo development take place inside the megasporangium. S. In these species. but the developing female gametophyte does not come out of the spore wall.. Some species of Selaginella (e. Development of embryo . rupestris. The sporangium is shed after the development of root and primary shoot of the new sporophyte. antherozoids swim in rain or dew water and reach the archegonia. when shed from the microsporangium (present in the distal part of the strobilus) lands on the partially open megasporangium. both the male and the female gametophytes lie within the megasporangium. After liberation from the male gametophyte. at this stage. the sporangium has only a single megaspore and at maturity of the archegonium the spore wall ruptures. apoda) show seed habit. it has all typical characters of a seed.

This eventually functions as the apical cell of the embryonic shoot. The rest of the embryo develops from the embryonic cell. At first the foot grows on one side but eventually comes to lie opposite the suspensor. total four cells) divide transversely to form two tiers of four cells each. the suspensor cell repeatedly divides to form a suspensor.The diploid sporophytic zygote is generation. and thus a four-celled embryo is formed. and thus an apical cell with three cutting faces is established.The remaining three cells of the 4-celled embryo and the sister cell of the apical cell (i. As development proceeds. 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.e. establishing an epibasal (upper) suspensor cell and a hypobasal (lower) embryonic cell. The cells of both the tiers divide irregularly forming a multicellular embryo. The derivatives of the lower tier form the foot.. The foot acts as a haustorial . It divides by two vertical walls at right angles to each other. the mother cell of the It divides transversely. One of the four cells of the embryo divides by an oblique vertical wall. which pushes the developing embryo deep into the female gametophyte.

After the formation of cotyledons and stem. . Roots. develop at the apex of the rhizophore. called 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.organ. In the axil of each cotyledon a ligule develops. At this stage. which eventually forms a cotyledon. in fact. The stem grows with the help of the apical cell of the embryo. the apical cell of the root differentiates on the lateral surface of the foot. But after the establishment of root and stem. a superficial cell in each of the two diagonally opposed quadrants of the upper tier differentiates as the apical cell of a foliar appendage. its main function is to absorb nutrition for the developing sporophyte from the female gametophyte. The part of the embryo immediately posterior to cotyledons develops into hypocotyledonary part of the stem. The derivatives of this cell develop into a root-like structure. the sporophyte becomes independent.

Although most reports of the medicinal uses of Selaginella are anecdotal. In Columbia. researchers have begun to identify and characterize the active compounds in Selaginella extracts (Kang et . articulata is used to treat snakebites and neutralize Bothrops atrox venom. In India. 2005). S. bryopteris is referred to as Sanjeevani—one that infuses life—for its medicinal properties (Sah et al. Selaginella is used as a popular herb for the treatment of various ailments (Lin and Kan. S. Throughout southern China. Pan et al 2001 and Maa et al 2003). 1990.Fig 11: General life-cycle of Selaginella Medicinal uses Many species of Selaginella have been used as traditional medicines.

Chen et al. Among the best characterized are uncinoside A and uncinoside B. 2001. Yang et al. 2006). Other biflavonoids from S. moellendorffii selectively inhibits the growth of some cancer cells by inducing apoptosis (Sun et al 1997. which are involved in the pathogenesis of some cancers (Lala and Chakraborty. The biflavone ginkgetin from S. 2005 and Yin et al. . biflavonoids that have potent antiviral activities against respiratory syncytial virus (Ma et al 2003). Zha et al 2004). 2005). 2004. tamariscina inhibit the induction of nitric oxide (NO) and prostaglandins (Pokharel et al Woo et al 2006. Su et al 2000).