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Morphology of Marsilea

Marsilea, commonly known as water fern, is represented

by about 53 living and 10 fossil species. The living
species occur in all parts of the world, but are more
common in the warmer parts of the world, such as
tropical Africa and Australia. They are aquatic or
amphibious; the aquatic species usually grow in shallow
ponds, but fruiting bodies (sporocarps) are formed only
in the terrestrial habitats. The amphibious species grow
in water-logged soil, partly submerged. Marsilea
condensate and M. rajasthanensis are near xerophytic
and M. hirsuta, an Australian species, can withstand long
dry spells. In India, the genus is represented by 9 living
species; of these M. minuta is the most widely
distributed. It is practically found all over India. In
Punjab, it is very common after the rains. M. brachypus,
M. quadrifolia, M. rajasthanensis and M. aegyptiaca are
the other important Indian species. Four species have
been reported from Rajasthan. Species, such as M.
minuta and M. quadrifolia are hydrophytic. They
grow submerged or partially out of water.

Systematic position

Kingdom : Plantae

Division :
Pteridophyta

Sub-division : Pteropsida

Class : Filicophyta

Order : Marsileales

Family :
Marsileaceae

Genus : Marsilea
Linn.

Common Name : Water Fern

The plant body of Marsilea is sporophytic having

diploid chromosome number in its somatic cells. During
sexual reproduction, it produces male and female
gametophytes, which are haploid in nature and form the
sperm and egg, respectively. The sporophyte is
differentiated into rhizome, roots, and leaves.

Rhizome (Stem): The stem is a long and slender

rhizome (Fig. 1). It creeps either on the surface like
stolon, or slightly below the surface of the soil like
rhizome. It grows extensively and is branched. The
branches arise at the bases of petioles and are extra-
axillary in position, arising in the lateral or oblique
position. They run in all directions and may get rooted at
the nodes. In this way, a single plant may cover an
extensive area of about 20 metre’s diameter or even
more. The stem is divisible into distinct nodes and
internodes. The internodes are long in the hydrophytic
species, and short in the sub-terrestrial or xerophytic
species.
 Fig. 1 Creeping and slender
rhizome of Marsilea sp.

Root: The primary root formed on the stem is short-lived

and is soon replaced by adventitious roots, which arise
gradually at the nodes on the underside of the stem (Fig.
1). Sometimes, roots may also arise at internodes
(Marsilea aegyptiaca), or laterally (M. minuta). The roots
are thin and may be branched or unbranched. They
develop in an acropetal sequence, i.e. the youngest root
is towards the apex of the rhizome. The number of roots
at a node and their size vary considerably.

Leaves: The leaves arise from the upper side of the stem
and are arranged in two alternate rows (Fig. 1). They are
long-petioled and compound. The petioles of the
submerged plants are long, thin and flexible, with the
lamina floating on the surface of water, while the leaves
of plants growing in mud or on land have upright, short
petioles, with lamina held in a spreading position. The
lamina is usually divided into four leaflets of the same
size. They spring from the tip of the petiole, so that the
leaf apparently looks quadrifoliate. Occasionally, the
number of leaflets may be 5 or 6, or even 8, instead of
the usual 4. In outline the leaflets are obovate, elliptical,
or wedge-shaped, with entire or dentate margins.

According to Puri and Garg (1953), the leaf of

Marsilea is pinnately compound, with four pinnules borne
on the slender rachis; two pinnules are noticeably higher
than the other two, and are inserted on the rachis in
alternate fashion. A leaflet has many dichotomously
branched veins, which are joined with each other by
transverse bands and their ends unite to form marginal
loops (Fig. 2).
 Fig. 2 Dichotomously branched
and petiolate leaves of Marsilea sp.

At the base of the petiole many bean-shaped or oval

and stalked sporocarps develop (Fig. 3).
Fig. 3 Bean-shaped or oval and stalked sporocarps
at the base of petioles in Marsilea sp.

Anatomy of Marsilea
Rhizome (Stem)
 Epidermis is single-layered, made up of compactly
arranged thick-walled cells.

 Cortex is differentiated into three regions, viz. outer

cortex, the middle cortex and the inner cortex:

a. outer cortex is aerenchymatous, one to several

cells in thickness, sometimes also consists of
tannin cells,

b. middle cortex consists of sclerenchymatous

tissue filled with air cavities arranged in the
form of ring, and

c. inner cortex is solid tissue of several cells in

thickness. The inner layer of inner cortex is
 parenchymatous, filled with starch, while outer
region of inner cortex is sclerenchymatous in
nature.

 The vascular cylinder is siphonostele; limited

externally and internally by endodermis, hence called
outer endodermis and inner endodermis,
respectively.

 The siphonostele is medullated. The xylem is in the

form of ring. Phloem is present on both sides of
xylem. Such a stele is called amphiphloic
siphonostele.

 The deposition of the several tissues from inwards is

outer endodermis, outer pericycle, outer phloem,
xylem, inner phloem, inner pericycle and inner
endodermis (Fig. 4).
 Fig. 4 Marsilea: Transverse section of
rhizome; a part cellular

Petiole

 Single layered epidermis made up of rectangular

cells.

 The outer cortex consists of few layers of thin walled

cells; middle cortex is aeranchymatous consisting of
ring of air-chambers; while the inner cortex is a
solid, compact tissue several cells deep. The inner
layers of inner cortex are paranchymatous and filled
with starch. Here and there tannin-filled cells also
occur.
  The stele is covered externally by a single-layered
distinct endodermis.

 It is triangular in outline, lies in the centre, and has a

single vascular bundle. The xylem is ‘V’ shaped,
with opening towards the axis.

 The order of the tissues from inner to outer side of

the stele is xylem, xylem parenchyma, phloem,
pericycle and endodermis (Fig. 5).

Fig. 5 Marsilea:Transverse section of

petiole; a part cellular

Leaflet
 It consists of upper and lower epidermal layers, each
made up of a single layer of paranchymatous cells.
The continuity of epidermis is interrupted by slightly
sunken stomata. Stomata are restricted to the upper
epidermis in floating leaves.

 The ground tissue (mesophyll) lies between the

upper and lower epidermis and is differentiated into
palisade and spongy parenchyma:

The palisade part lies just beneath the upper

epidermis and consists of columnar cells rich in
chloroplasts, while the spongy part faces the lower
epidermis and consists of rounded cells

In case of submerged species, there is no

distinction between palisade and spongy tissues.

 Vascular bundles are embedded in the mesophyll

tissue. They are concentric in nature. Each bundle
has a central core of xylem, surrounded by phloem.

 The arrangement of tissues in the stele is xylem,

phloem and bundle sheath (Fig. 6).
 Fig. 6 Marsilea: Transverse
section of lamina.

Root

 It consists of piliferous layer instead of epidermis,

made up of compactly arranged biconvex cells.

 Beneath piliferous layer, the cortex is made up of an

outer cortex and inner cortex:

Outer cortex is aerenchymatous, consisting of

large air chambers arranged in the form of ring.

Inner cortex is compact, made up of rounded

cells containing starch.
 Inner cortex is delimited by a single layer of distinct
endodermis.

 Vascular bundle lies within the endodermis, and is

usually diarch and exarch.

 The xylem is plate-like and occupies the centre of

stele. The protoxylem, consisting of two small mass
of cells, is towards the periphery (pericycle), while
metaxylem is large, plate-like and occupies the
centre.

 There is no medullary tissue (pith) in the stele and

arrangement of tissue is metaxylem, protoxylem,
phloem, pericycle and endodermis (Fig. 7).
 Fig. 7 Marsilea: Transverse section of
root.

Reproduction in Marsilea

Marsilea is a sporophyte and reproduces asexually both

by vegetative means as well as by means of spores.

Vegetative reproduction: In Marsilea, vegetative

reproduction occurs by the formation of tubers, which are
small, bud-like structures containing reserve food
material, arising from some subterranean branches of the
rhizome. These tubers serve as perennating organs, and
are capable of tiding over the unfavourable conditions.
On the return of favourable conditions, these tubers
germinate to form new plants. Tuber formation has been
reported in a few Marsilea species, such as M. minuta and
M. hirsuta.

Reproduction by spores (spore formation): Marsilea

is a heterosporous fern, which produces two types of
spores – microspores and megaspores in separate
sporangia, borne in special bean-shaped bodies called the
sporocarps.
Sporocarps: Sporocarp is a bean-shaped to ovoid,
nutlike structure, attached to the basal part of the petiole
with the help of a stalk. It is green and soft when young,
but turns dark-brown at maturity. Usually one sporocarp
is present at the base of each petiole, but in some
species, the number varies from 2 – 20. Sometimes, the
attachment of sporocarps with the petiole shows so much
variation that different species can be distinguished on
this particular character (Fig. 8); for example in M.
polycarpa many sporocarps are attached on one side of
the petiole in a single vertical row. In M. quadrifolia,
pedicels (stalks) are united with one another, and then
jointly inserted on the petiole. In M. minuta, the stalks of
all the sporocarps though free, are attached to the petiole
at a single point.
 Fig. 8 Marsilea: different modes of attachment of
sporocarps to the petiole – A, M.
polycarpa; B, M. quadrifolia; C, M. minuta; and D, M.
vestita.
Structure of sporocarp: The sporocarp is differentiated
into a stalk (pedicel) and a body. The stalk is fused
laterally to the back of the body of the sporocarp,
generally forming a distinct ridge called ‘raphe’. In some
species, the distal (upper) end of the raphe is marked by
one or two teeth-like projections, known as tubercles.

Internal structure of the sporocarp: The internal

structure of the sporocarp can be studied under the
following headings:

Sporocarp wall: The wall of the sporocarp is very hard,

thick and highly resistant to mechanical injury. It is
differentiated into an outer epidermis, a middle
hypodermis, and an inner paranchymatous zone. The
epidermis is made up of cuboidal cells, covered with a
thick layer of cuticle. A large number of sunken stomata
are present in the epidermis. The hypodermis consists of
two layers of radially-elongated palisade-like cells, which
are compactly arranged without any intercellular spaces
between them. The inner paranchymatous cells of this
zone form a gelatinous ring inside the sporocarp wall
(Fig. 9).

Fig. 9 Transverse section of the wall of Marsilea

sporocarp: A, young stage; B, old stage.

Sori: When we cut the horizontal section of sporocarp, a

ring appears in the form of a dorsal and a ventral mass.
In this plane, both micro- and megasporangia are visible
as sori. The sori are the reproductive structures arranged
in the two alternating rows in the cavity of the sporocarp
(Fig. 10). Each sorus has a receptacle which contains one
terminal megasporangium and two microsporangia on the
lateral sides. It is surrounded by an indusium. The sori
overlap each other and the indusia of adjacent sori are
partially fused.
Fig. 10 Marsilea sp.: A, Vertical transverse section
of sporocarp; B, horizontal longitudinal section of
sporocarp.

The number of sori in a sporocarp varies from two in M.

aegyptica to twenty in M. quadrifolia and M. vestita.
There are 11 – 12 sori in M. minuta. Each sorus bears
both mega and microsporangia. The former are short-
stalked and are arranged in a row at the tip of the
receptacle, whereas the latter are long-stalked and arise
on the sides. The number of micro- and megasporangia
varies with species. In M. minuta, a sorus has 4-8
megasporangia and 8-13 microsporangia.
Development of sporocarp: Sporocarp originates from
one of the marginal cells of a very young leaf, when it is
6-8 cells high. One of the marginal cell cuts off segments
on either side and one of the derivatives acts as the
apical cell for the second sporocarp. This process is
repeated several times depending on the number of
sporocarps formed at the base of a leaf. The sporocarp,
initially cuts off segments on either side and forms a
mass of undifferentiated cells, which later curves
markedly so that its distal end is directed in a horizontal
plane. Subsequently, two rows of soral mother cells are
differentiated on the ventral side of the young sporocarp.
The differential growth of soral mother cells and their
derivatives, results in the formation of involutions on the
upper surface of the sorus. These involutions grow into
arc-shaped depressions called soral canals. The soral
mother cells divide to form receptacular surface in the
alternating rows. The sporangial initial develops on the
receptacular surface in acropetal succession, i.e. first
sporangial initial is formed at the apex of receptacle and
it forms megasporangia. Subsequently, two initials are
formed on the lateral sides of the receptacle. They
functions as microsporangial initial, each forming a
microsporangium (Fig. 11).

Fig. 11 Diagrams showing developmental stages of

micro- and megasporangia of Marsilea sp.
Development of sporangia: Since Marsilea is a
heterosporous fern, megasporangia (macrosporangia)
produce megaspores and the microsporangia produce
microspores. The sporangia develop in a basipetalous
manner on the receptacle. The megasporangia develop
first at the top of the receptacle and are, therefore, older,
while the microsporangia develop later along the sides of
the receptacle. The development of microsporangium, as
well as of megasporangium is almost similar. The
sporangial initial divides transversely, forming an outer
and an inner cell. The entire sporangium develops from
the outer cell, the inner cell taking no part in its
development. The outer cell of sporangium, which is
destined to form spore mother cell, undergoes repeated
divisions. Each spore mother cell undergoes reduction
division and forms four haploid spores. In
megasporangia, all but one spore degenerates to form a
nutritive liquid, which grows in size at the expense of
others, forming a single large functional megaspore (Fig.
12). In microsporangia, almost all the spores in a
microsporangium are functional. They are smaller in size.
Fig. 12 Marsilea sp.: mature megaspore within the
megasporangium, showing very thick and well-
differentiated wall.

Dehiscence of sporocarp: There is no special

mechanism for the liberation of sporangia from the
sporocarp. They are set free only when the wall of the
sporocarp splits open. As the sporocarp breaks at one
point due to absorption of water, the sori come out along
the gelatinous ring of sporocarp (Fig. 13). These sori
function as mother cells of the gematophytic generation.
 Fig. 13 Diagrams showing stages in the
dehiscence of Marsilea sporocarps.

Gematophytic generation

The microspores and the megaspores germinate

separately to produce male (micro-) and the female
(mega-) gametophytes. The microspores germinate
immediately after shedding, forming small biconvex
prothallial cell and a large apical cell. The apical cell by
repeated divisions ultimately forms androcytes or
antherizoid- mother cells. Each androcyte
metamorphoses into a cork screw-shaped multiflagellate
antherizoid, characterized by the presence of a prominent
terminal vesicle (Fig. 14).
 Fig. 14 Marsilea: screw-shaped multiflagellate
antherizoid, characterized by the
presence of a prominent terminal
vesicle.

Similarly, megaspore undergoes division, forming

small nipple-shaped apical cell and a large basal
prothallial cell. The apical cell forms the female
gametophyte, while prothallial cell provides nutrition to
the developing gematophytes (Fig. 15).
 Fig. 15 Marsilea: germination and
development of the female gametophyte.

Fertilization: The neck canal cells and the ventral canal

cell degenerates to form a mucilaginous mass, which
attracts sperms chemotactically. One of the sperms fuses
with the egg to accomplish fertilization and form the
diploid zygote. The zygote is the mother cell of the next
sporophytic generation and develops into a young
sporophyte within 2-4 days after fertilization.

Embryogeny: The fertilized egg (zygote) enlarges in its

size and secretes a thin cellulose wall around it. It enters
into rapid divisions, to give rise to a young sporophyte
within 2 - 4 days. Zygote divides vertically and produces
two unequal cells. The larger cell is known as the
epibasal cell, and the smaller one as the hypobasal cell.
These two cells undergo transverse divisions to form a
quadrant embryo. The cells of the quadrant embryo show
definite relation to the primary growth of the embryo.
Epibasal half produces shoot and leaf. Hypobasal half
produces root and foot. Hence the development is
described as lateral. In quadrant embryo, the next
divisions are irregular. At the time of differentiation of
different primordia in the embryo, the adjoining cells of
megagametophyte divide periclinally, forming a thick
calyptras, which encloses and protects the developing
embryo. The calyptra is greenish in colour. Large number
of rhizoids may develop from the base of calyptra. The
embryo gives rise to the young seedling that penetrates
through the calyptra and grows into a sporophyte (Fig.
16).
Fig. 16 Marsilea: stages in the development of
embryo - A-E, early developmental stages; F,
mature female gametophyte bearing young embryo
surrounded by the calyptra; G, median-longitudinal
section of the mature female gametophyte.

Life cycle: In the life cycle of Marsilea, we find

alternation of generations. Marsilea plant is a sporophyte,
which is the dominant phase in the entire life cycle. The
sporophyte comprises rhizome, roots and leaves. It
reproduces vegetatively, as well as by means of spores.
The spores are formed in sporangia on specialized
structures called sporocarps. The sporocarps bear two
kinds of asexual spores which are formed after meiosis
and are, thus haploid (meiospores). The smaller spores
are known as microspores. They are produced within
microsporangia. The larger spores are known as
megaspores, which are formed in the megasporangia.
Both the sporangia are produced within the fruit body,
the sporocarp. The whole life cycle of Marsilea is depicted
in Fig. 17.

Fig. 17 Marsilea: diagrammatic

representation of life cycle.