Pteropsida

Distinguishing features

1. The sporophytic plant body is differentiated into roots, stem and spirally arranged leaves.
2. Most of the members grow in moist and shady habitats, with their luxuriant growth in
humid forests of tropical regions.
3. The stele in Pteropsida shows a wide variety of types. It varies from simple protostelic type
to complex siphonostelic, solenostelic or dictyostelic conditions.
4. The vegetative propagation takes place by a variety of ways, like fragmentation (e.g.
Pteridium, Dryopteris, Adiantum, Pteris, etc.), adventitious buds (e.g. Asplenium bulbiferum,
Cystopteris bulbifera, etc.), stem tubers (Marsilea) and also by apogamy (e.g. Marsilea and
many other ferns).
5. The spores, formed in sporangia, are thin-walled or thick-walled. Members are either
homosporous or heterosporous. The sporangia occur in groups called sori. The sori are either
arranged on the margins or abaxial surface of leaves or leaflets. Sometimes, they are borne
terminally on the axis. The sporangia-bearing leaves are called sporophylls. The sori may be
unprotected or naked, or they may be protected by the revoluted margins of the sporophyll or
by a special outgrowth called indusium. In genera like Adiantum and Pteris, a false indusium
is present. The spores germinate and give rise to prothalli.
6. The antheridia may be embedded or emergent, and contain many multiflagellate
antherozoids.
Systematic position

Class: Pteropsida
Subclass: Leptosporangiatae
Order: Filicales
Family: Polypodiaceae
Genus: Pteris
 Pteris sp. - Occurrence and Distribution:
• This is a cosmopolitan fern being distributed
in almost all geographical regions. Pteris
however, prefers tropical and sub-tropical
climates. Plants usually grow in well drained
places or in the crevices of rocks. They are
very common along the slopes of hills and can
be seen even at 1200 metres above sea level.

• There are about 250-280 species reported for

the genus. Some of the common Indian species
are P. quadriaurita, P.critica, P.vittata,
P.pellucida, P.wallichiana, P.stenophylla,
P.biaurita, etc. Tanaka and Chin-ming (1982)
have isolated Pterosin and Astragalin from
Pteridium acquilinum sub spp Wightianum.

Creeping rhizome/ rhizomatous stem, covered with scale, hair are absent,
Roots arise from the lower surface, leaves arise from the upper surface.
Leaves are pinnately compound looks as frond, petiole is covered with scale,
Pinnae are smaller at the base. Larger near the middle, smaller towards the apex.
Pteris : Structure of Sporophyte
Sporophyte:
Morphology of the Plant:
• The plant body has a rhizomatous stem that
produces roots and leaves. The rhizome may be
creeping (P.grandiflora) or compact and erect as in
P.cretica and P.longifolia. The rhizome may or may
not show branching and is covered with scales.
Roots arise at the base of the leaf or all over the
rhizome. The growing point of the rhizome is
covered with ramenta.
• The leaves arise from the upper surface of the
rhizome and have a long rachis. They are
unipinnately compound, decompound or multi-
pinnately compound. The dissections of the pinnae
are not as deep as in Pteridium.
• Venation is of open furcate type. The pinnae are
small near the base, large towards the middle and
once again small towards the apex as in P. vittata.
The leaves show circinate vernation that is typical
to true ferns.
• Very often pinnae are Coriaceous. The leaves bear
reproductive structures-sorus along the ventral
margin of pinnae. The sorus is continuous along the
margin but avoiding the apices of the segments and
usually in the sinuses between them.
Internal Structure of Sporophyte
Rhizome:
• The rhizome in a transection exhibits quite a good amount of
diversity particularly in the stelar organisation. It is
solenostelic (P. grandiflora, P.vittata), or dictyostelic. In P.
biaurita the rhizome shows a mixed protostele in the lower
region. It becomes siphonostelic a little up, and near the apex
it shows a polycyclic dictyostelic organisation.

• The epidermis is single layered and it encloses a broad cortex.

There is a hypodermal sclerenchyma. The bulk of the cortex
however, is made up of parenchyma. The stele consists of a
number of meristeles. Usually there are two rings of
meristeles. The inner ring consists of 2 or 3 large meristeles
whereas the outer ring consists of a number of small
meristeles.

• Each meristele has a band or plate like xylem mass.

Sometimes it is angular. One or two protoxylem groups are
embedded in the metaxylem (mesarch). Surrounding the
xylem is the phloem. Each meristele may have its own
endodermis. Here the breaking of the vascular strand is due to
the leaf gaps.
Internal Structure of Sporophyte
Leaf:
• The leaves are borne on the upper surface of the rhizome.
When young the leaves are spirally coiled and show
circinate vernation that is typical of true ferns. The leaves
are unipinnately or multipinnately compound or
decompound with a long rachis.
• The pinnae are small near the base as well as towards the
apex, while they are large towards the middle. The pinnae
are very often coriaceous. All leaves are fertile, bearing
sori along the ventral margin of pinnae, except the apices
of the segments.
• The rachis is traversed by a single C/U/ V-shaped leaf
trace. The lamina is bifacial, hypostomatic. Mesophyll
cells may or may not be differentiated. A concentric
vascular bundle with distinct bundle sheath is present in
the midrib.
• The rachis is traversed by a single leaf trace which is
variable in shape. ‘C’ shaped leaf traces are found in P.
vittata. In P. biaurita the leaf trace is ‘U’ shaped while
entering the leaf base, but further up it becomes ‘V’
shaped.
• The xylem strand appears hooked. As usual xylem is
surrounded by phloem, pericycle and endodermis. The
cortical region has an inner parenchymatous zone and an
outer sclerotic zone. Epidermis is single layered with a
deposition the cuticle over it. Ramenta arise from some of
the epidermal cells.
• The lamina appears bifacial. It may or may not have a
differentiated mesophyll. Leaves are generally
hypostomatic. The midrib has a concentric vascular bundle
(P. vittata) with a distinct endodermis (bundle sheath).
Reproduction in Pteris

Pteris reproduces by means of spores.

Spore-Producing Organ:
• Pteris is a homosporous fern. The sorus of
Pteris is called coenosorus. Coenosori are
marginal, borne continuously on a vascular
commissure connected with vein ends. Thus
the sporangia of Pteris form a continuous
linear sorus along the margin, hence the indivi-
duality of sori is lost.
• The coenosori are protected by the reflexed
margin (false indusium) of the pinnae. Sori are
of mixed type intermingled with many sterile
hairs in between the sporangia.
Structure of a Mature Sporangium
• A mature sporangium has a long stalk that terminates
in a capsule
• The jacket of the capsule is single-layered, but with
three different types of cells
• (1)A thick walled vertical annulus incompletely
overarches the sporangium,
• (2) A thin-walled radially arranged stomium, and
• (3) Large parenchymatous cells with undulated
walls.
• The capsule contains 48 spores. All spores are
structurally and functionally alike; hence Pteris is a
homosporous pteridophyte. Spores are triangular in
shape with trilete aperture, bounded by two walls.
The outer wall, exine, is variously ornamented.
• The sporangium dehisces transversely along the
stomium due to the shrinkage of annular cells. The
spores are dispersed through air to a moderate
distance.
Structure of Sporangium
Development of Sporangium
• The development of sporangium in Pteris is of
leptosporangiate type. A single superficial cell of the
receptacle functions as the sporangial initial which divides
transversely to produce an upper cell and a lower cell.

• The lower cell does not take part in sporangium

development, while the upper cell, by intersecting oblique
walls, gets differentiated into an apical cell with three
cutting faces. The apical cell cuts off two segments along
each of its three cutting faces.

• The apical cell divides periclinally to form an outer jacket

initial and an inner tetrahedral archesporial cell. The jacket
initial divides, anticlinally to form a single-layered jacket
of the sporangium. The archesporial cell further divides
periclinally to form an outer tapetal initial and an inner
primary sporogenous cell.

• The tapetal initial by one periclinal and several anticlinal

divisions forms two-layered tapetum. The primary
sporogenous cell divides to form 12 spore mother cells.
The spore mother cells divide meiotically to produce
haploid spores, while the tapetal cells disorganise and
provide nutrition to the spores.
Dehiscence of Sporangium
Structure of Gametophyte
• The spores germinate after falling on a suitable substratum.
Initially the spore wall (exine) ruptures and the inner
contents come out in the form of a germ tube and
subsequently by a transverse division in the germ tube forms
the first rhizoid and the first prothallial cell. The prothallial
cell divides to form a small filament having an apical
terminal cell with two cutting faces.

• The apical cell further divides and a thin prothallus is

formed first. Finally a mature prothallus is formed which
becomes cordate, dorsiventrally flattened, aerial and
photosynthetic.

• The prothallus is made up of parenchymatous cells which

are single-celled thick towards the margin and many-celled
thick towards the centre. The growing point are located in
the apical notch. Rhizoids are formed over the ventral
surface.

• The prothallus is monoecious, protandrous. Antheridia

appear first and are confined to the basal central or lateral
regions among the rhizoids. Archegonia develop near the
apical notch..
Development and Structure of the Antheridium
One of the superficial cells on the prothallus
towards the ventral surface projects out a little
and functions as an antheridial initial. This
undergoes a transverse division to form two
superposed cells.
Due to the increase of turgor pressure in the
upper cell, the cross wall bulges down and
ultimately touches the wall of the lower cell.
The upper cell now divides to form a dome cell
and the primary androgonial cell.
• At this stage the young antheridium consists of
a dome cell, a primary androgonial cell and a
ring cell (lower cell looking like a ring). The
dome cell divides transversely to form a cap
cell and the second ring cell.
• The cap cell may are may not divide to form
two cover cells. Meanwhile the primary
androgonial cell divides to form 20-25
androcytes.
A mature antheridium has two ring cells, one or
two cover cells enclosing a mass of
antherozoids which are coiled and multi-ciliate.
At maturity the antheridium absorbs water and
swells, the cover cell opens out releasing the
antherozoids.
Development and Structure of the Archegonium:
Archegonium:
• The archegonium is also initiated in the derivates of apical
meristem. One of the superficial cells functions as
archegonial initial which, on periclinal division, forms an
outer primary cover cell and an inner cell. The primary
cover cell, by two vertical divisions at right angles to each
other, forms quadrant of neck initials.
• Further anticlinal divisions of neck initials form the neck
of the archegonium. The neck barely protrudes out of the
thallus. The inner cell, on the other hand, divides
periclinally (transverse) to form a basal cell and a central
cell.
• The central cell divides transversely into a primary canal
cell and a primary venter cell. The primary canal cell
directly functions as neck canal cell. The primary venter
cell, however, divides transversely and forms a ventral
canal cell and a large egg.
• At maturity, the ventral canal cell, the neck canal cell and
the neck cells at the top are well- disorganised, they thus
form an open passage for the antherozoids to come
towards the egg
• A mature archegonium of Pteris consists of a 5-6 celled
projecting curved neck, a neck canal cell, a ventral canal
cell and an egg.
.
Development of New Sporophyte
Fertilisation:
The antheridium at maturity absorbs water and swells.
Due to the increase in pressure within the antheridium
the cover cells split apart releasing the antherozoids in
a thin film of water present on the surface of the
prothallus.
• At the same time the ventral canal cell, the neck canal
cell and the neck cells at the top disintegrate forming
an open passage for the antherozoids to come towards
the egg and, eventually, one of the antherozoids fuses
with the egg to form the zygote
• New Sporophyte (Embryo):
• Like other leptosporangiate ferns, in Pteris the first
division of the zygote is vertical, followed by a second
transverse division resulting in the formation of a
quadrant. Further a 32-celled embryo is formed due to
further divisions of the quadrant.
• The differentiation of embryo begins at this 32-celled
stage. No suspensor is formed; the hypobasal cells
form stem apex and foot, while epibasal cells form
cotyledon and root. With the development of embryo,
the venter of the archegonium forms a protective layer,
called catyptra, around the embryo.
• In the young embryo the root and cotyledon grow more
rapidly than the shoot. The root pierces the prothallus
and establishes the sporeling in the soil. Later, the first
leaf develops.
Life Cycle of Pteris