________PTERIDOPHYTES__________

The term 'Pteridophyta' is evolved from the Greek word, 'Pteron' means feather, as most of
the pteridophytes show feather like (pinnately compound) leaves. Pteridophytes are considered to
be the first cryptogams (seedless plants) which have successfully invaded the land. They differ
from other cryptogams mainly in having well developed vascular tissue system; hence they are
also called vascular cryptogams.
The living pteridophytes are mainly herbaceous except a few woody tree ferns. They exhibit
great variation in form, size and structure and show the following distinguishing characters:
i) The life history shows two distinct stages, viz., spore producing stage called sporophyte
and gametes producing stage called gametophyte.
ii) Both sporophyte and gametophyte are independent stages.
iii) Sportophyte is the dominant stage distinguished into roots, stem and leaves and shows well
developed vascular tissue with xylem and phloem well differentiated.
iv) The gametophyte is insignificant but independent stage. Sex organs are
multicellular, jacketed but without stalk.
v) In life-history, both the stages regularly alternate with each other and show distinct
alternation of generations.
The various classification system were proposed by different authorities during the 20 th
Century. However, classification followed in this book is as given by G. M. Smith
(1955).___________________________________________________________________________
~________________________________NEPHROLEPIS__________________________________
Division : Pterophyta i) Sporophyte differentiated into roots, stem and leaves.

ii) Sporangia produced on leaves, either along the

margin or on abaxial side.
iii) Large pinnately compound leaves.
iv) Gametophyte is small, green and free-living.
Class - Filicinae Characters same as above (Only class of the division)

Sub-class i Leptosporangiatae i) Sporangium develops from single initial cell.

ii) Jacket of the sporangium is one cell in
thickness.
iii) Sporangia are not laterally united to form
synangium.
Order : Filicales i) Sori are simple, gradate or mixed.
 «) All the spores are of same type, hence son are
homosporous.
Family ; Polypodiaceae i) Sori are of mixed type.
ii) Sporangium has vertical annul us and
transverse dehiscence, with stomhim.
iii) Sporangium contains 32 to 64 spor=
iv) In the early stages the game is
filamentous and at maturity becomes heart-
shaped.
v) Jacket of antheridium has 3 cells. Antherozoids
are relatively few in number.
Genus : Nephrolepis i) Spores with kidney-shaped indusium.
ii) Each sporangium contains 64 spores.
Occurrence
Ferns are highly developed vascular cryptogams belonging to the large group
pteridophyta. They have worldwide distribution showing diversity in habits and habitats.
However, they are usually moisture and shade loving plants abundant in tropical rain forests.
They exhibit many beautiful varieties, which are often cultivated in garden as ornamental
plants. The common Indian pteridophytic species, Nephrolqiis exaltata, is described below:
Sporophyte
External Morphology
Sporophyte is the dominant stage and it is
differentiated into roots, stem and leaves with
true vasculature (conductive strands).
Roots
Roots arise from base of the stem and
stolon forming adventitious fibrous root
system. They show persistent root hair. They
absorb mineral salts and water and fix the
plant in the soil.
Stem
Stem is about two inches in height, thick,
woody, brown in colour. It is called rhizomatic
stem or stock or caudex because it is obliquely
placed in the soil and partially underground. It
produces stolons and compound leaves from its
aerial parts and adventitious fibrous roots from the
underground part. It is rough due to persistent leaf
bases.
Stolons
Position : Stolons arise from rhizomatic stem
and are extra-axillary in position.
Structure : These are wire-like, thin
branches which are green when young and turn
brown, at maturity. Each stolon grows slightlv
upwards and bends downwards to form an arch in
the soil. At the touching point, stolon produces
adventitious roots below and bud or spur above,
which develops into new sporophyte.
Function : As the stolon gives rise to new
sporophytes. it performs the function of vegetative reproduction.
Ramenta
Position : These are
brown scale-like epidermal
outgrowths densely covering
the rhizome, stolons, petiole
and rachis.
Structure : Each
ramentum is multicellular,
membranous, more or less
triangular structure which is
single cell in thickness. The
cells are dry with thick brown
walls. The marginal cell gives
out hair like projections. The
ramentum is attached to the
epidermis at the broader end
by a dark brown point of
attachment.
Function : Ramenta are protective in function, guarding the plant from excessive heat and
rain water. They also retain water in their axils to provide humidity.
Leaves
Position : Leaves are spirally arranged on the aerial parts of the stem.
Structure : Each leaf is pinnately compound and shows central axis differentiated into
rachis and petiole (stipe). On rachis, leaflets are arranged on either side to form frond. Each
leaflet is almost sessile and show's lobed or auricled base. Upper auricle is large than the lower
one. The leaflet is lanceolate in shape, with crenate margin and acute apex. There is single midrib
but lateral veins are bifurcated at the tip to form forked venation. The leaflet shows presence of
hydathodes on the adaxial surface at each vein ending and sori on abaxial surface at the upper
vein ending of each bifurcated vein. The young leaves are silvery white in colour and are coiled
from apex downward to form circinate ptyxis.
Function : Leaf performs following functions :
i) It performs the
function of
photosynthesis.
ii) It removes
excess of water by
process of guttation
and transpiration.
iii) It
performs
the
function
of asexual
reproduction by producing sori.
Hydathodes
 Position: These are
white dots arranged in
two longitudinal rows
on adaxial (dorsal)
surface along the
margins of each pinna at
either vein ending of
each bifurcated vein.
Structure : It
consists of a group of small thin walled cells sunken into a minute oval depression and bounded
by closely placed two to three
hgpets erf rectangular epithelial cells which are, in 'turn, surrounded by normal epidermal
oefls.
Function : The glandular cell exude water containing calcium salts, by the process of
s_ttation (excretion in the form of liquid from plant under root pressure). It is due to :r:err,al
pressure developed as a result of active absorption and low transpiration. The cell; of hydathode
are hygroscopic and at times help to absorb moisture from the azr.osphere. They are called water
stomata, as they exude water in ice form of liquid and ~;: in the form of water vapour unlike
normal stoma. When water evaporates from the surface of hydathode, calcium carbonate is
precipitated as white dots, hence, hydathodes are sometimes also known as chalk glands.
Transpiration Guttation
1 Takes place mainly through stomata. 1. Takes place through hydathodes.
2. Water is given out in the form of water 2. Water is given out in the form of water
vapour. droplets.
3. Takes place under transpiration pull. 3. Takes place under root pressure.
4. Controlled by closing and opening of guard 4. Guard cells are absent.
cells.
5. Ca salts are not given out. 5. Calcium salts are given out.
______________
Internal Morphology T.S. of rachis
In general outline, the section appears circular with a shallow groove on adaxial (dorsal)
side. It is differentiated into epidermis, ground tissue and vascular system.
Epidermis consists
of single layer of closely
packed rectangular cell
,with their outer walls
cutinized.
Ground tissue is
differentiated into the
outer zone of thick,
walled lignified cells
forming hypodermis and
the rest of inner ground
tissue which
is
1
parenchymatous. The
cells of the ground tissue
contain starch grains.
The outer cells may show
chloroplasts

when
young. The layer of the
ground

tissue
immediatel
y
Ii surrounding
each
vascular strand shows
cell with their inner and radial walls highly thickened due to phlobophaence forming a
distinct brown ring. "
Vascular system includes 2 to 5 vascular strands embedded in the ground tissue. The, are
arranged in a horse shoe-shaped manner which opens towards the groove. Each vascular stand
contains a single vascular bundle surrounded by its own single layered, thin wailed pericycle and
endodermis. Hence, each vascular strand is called meristele and the entire vascular system is
called dictyostele.
 Each vascular bundle is conjoint, concentric and amphicribral or hardocentric as
xylem is surrounded by phloem. Xylem is exarch, usually comma shaped or "C" shaped
with metaxylem at one end and protoxylem at the other end. The xylem consists of only
xylem tracheids and xylem parenchyma; vessels being absent. Phloem consists of only
sieve cell and phloem parenchyma. Sieve tubes and companion cell are absent.
T.S. of stolon
In general outline> the
section appears circular
and is distinguished into
epidermis, cortex and
single stele (monostelic).
Epidermis consists of
single layer of closely,
packed rectangular cell
with their outer walls
cutinised.
Cortex is
differentiated
into three regions, viz.,
outer, middle and inner.
Outer cortex is made up of
a
few layers of lignified cell
forming

hypodermis.
Middle cortex
is
parenchymatous.

The
 innermost layer of the
cortex is called
endodermis which is
made up of thin-walled,
recetangular,
parenchymatous cells. A
few layers of cells of the
cortex outside the
endodermis may become
thick-walled and brown.
Stele is single and
protostelic as it consists of
solid column of vascular
tissue without pith. It is
externally surrounding by
pericycle made up of two to three layers of thin walled cells. It includes a single vascular
bundle, which is conjoint, concentric, and hadrocentric (amphicribral) as xylem is
surrounded by phloem. The xylem is star-shaped with metaxylem in the centre and the
protoxylem radiating towards the periphery. Such a type of protostele with stellate (star-
shaped) xylem is called aetinostele. The xylem consists of only tracheids and xylem
parenchyma; vessels being absent. The phloem includes phloem parenchyma and sieve
cells which completely surround the xylem and become more prominent in-between the
radiating protoxylem groups. In phloem, sieve tubes and companion cells are absent.
T.S. of pinna passing through sorus
T.S. of pinna shows upper epidermis, mesophyll, lower epidermis and sorus.
Upper epidermis consists of single layer of closely packed, brick-shaped,
chloroplasts containing cells with their outer walls cutinized. Epidermis is distinguished
by the absence of stomata and presence of hydathode as shallow depression above vein
endings.
 Mesophyll consists of spongy tissue containing one type of cells (parenchymatous) with
chloroplasts; hence, the pinna is said to be isobilateral. However, the upper cells are more
compact than lower ones. It also shows vein endings in form of group of tracheids below
hydathode.
Lower epidermis is just like upper epidermis but with stoma and without hydathodes.
Sorus is attached to the lower epidermis at the vein ending and consists of:
i) Placenta, which is cushion-like parenchymatous structure providing nourishment to
the developing sporangia.
ii) Sporangia are present on either side of the placenta. Each sporangium consists of a
stalk and a capsule containing spores.
iii) Indusium, which is single layered, external protective covering. In T.S., it looks
like a mustache; however, in surface view, it appears kidney-shaped. It is attached
to the placenta by a short, massive, parenchymatous stalk.
Asexual reproduction
The asexual reproduction takes by
formation of sori, which are produced
abaxially on the leaflets. The leaf
bearing leaflets with sori is called
sporophyll.
a) Sorus

Position: Sori are present on abaxial

surface of leaflet, near both the margins at
upper vein endings of each bifurcated vein.
Structure : Each sorus is an asexual reproductive body. It consists of soft, cushion-like
parenchymatous tissue called placenta at the base on which sporangia are borne. They are
covered by kidney-shaped , sterile covering called indusium. Nephros means kidney, hence the
name Nqjhrolepis. The sorus is attached to leaflet by placenta.
 b) Sporangium

Definition : Sporangium or spore sac is

an asexual reproductive organ containing
diploid spore mother cells, which at maturity
undergo meiosis to form haploid spores.
Position : Sporangia are produced in
group on placenta and are covered by
indusium. The entire structure thus formed is
called sorus.
Structure : Each sporangium consists of basal stalk and distal capsule. The stalk is solid,
multicellular with cell arranged in two rows. The capsule is biconvex and consists of a capsule-wall
known as jacket, which is single layered and thin walled. Along the 3/4* edge, it shows cells of
annulus, with their inner tangential and radial walls are
thickened. At the remaining edge, there are transversely elongated thin walled cells.
Interior of the capsule is filled up with sixteen diploid spore mother cells, which at
maturity undergo meiosis to produce 64 haploid, kidney-shaped spores. As all the spores
are of same size and shape, the plant is homosporous.
Each spore is more or less kidney-shaped, slightly flattened, dark brown body. It is haploid
containing uninucleate protoplast with some reserve food and is covered with two coats, viz.,
exosporium, which is outer, thick, brown, resistive coat and endosporium, which is inner, thin coat.
c) Dehiscence of sporangium
 When the sporangium
matures and the weather is
dry; the dissemination of
spores takes place. In dry
weather, the outer thin walls
of the cells of annulus lose
water and shrink, exerting a
pressure on the transversely elongated thin-walled cells on one side. As a result of this, the
sporangium breaks open in-between these thin-walled lip-like cells, forming a transverse opening
called stomium. As the annulus loses more and more water, it shrinks further and goes backwards,
tearing the wall of capsule up to the annulus and carrying most of the spores along with it. At one
stage, when pressure on annulus is released, it act like a spring and suddenly snaps forward to its
original position so that the spores are violently thrown out, in the air to get carried away by wind.
Thus, the spores are liberated and dispersed.
 d) Germination of spore
During favorable conditions, the spore
germinates. The exosporium breaks and the
endosporium grow into a short germ tube.
The latter soon divides into two unequal
cells, viz., the smaller colourless rhizoidal
cell, which gives rise to unicellular rhizoids
and the other green prothallial cell. The
latter by repeated transverse cell divisions,
gives rise to green alga-like, uniseriate,
filamentous structure. Soon the divisions take place in other planes to form green plate-like body.
Owing to more rapid divisions and growth of the marginal cells; the plate-like structure eventually
gets converted into more or less heart-shaped structure called prothallus, which represents the
gametophytic generation of fern.
Gametophyte
The gametophyte of fern is
called prothallus. It is green,
delicate, dorsiventrally flattened,
heart-shaped body, measuring
about 5mm across. At the apex, it
shows a notch
and ventraily, at the narrow
posterior end, it bears unicellular
rhizoids helping in anchorage and
absorption of water and mineral
salts from the soil. The prothallus
is one-celled thick except towards
the notch where it is many layered
forming a cushion-like structure.
The cells of the prothallus are thin
walled and with full of
chloroplasts. There are neither
stomata nor intercellular spaces;
hence, exchange of gases takes
place directly.

Nutrition of prothallus is autotrophic as chlorenchymatous cells prepare food material by

photosynthesis. The rhizoids absorb mineral salts and water. Thus, prothallus is also an autotrophic,
independent generation.
Prothallus is monoecious as it bears both male and female sex organs i.e. antheridia and
archegonia respectively, on the same plant. They are present on the ventral surface. The antheridia
are present near the rhizoids while archegonia are present below the apical notch partially embedded
in the cushion.
Sexual Reproduction
Sexual reproduction is oogamous. It takes place by formation of well developed, multicellular,
jacketed sex organ.
a) Sex Organs : Prothallus is monoecious as it bears both male and female sex organs i.e.
antheridia and archegonia on the same plant. They are present on the ventral surface.
Antheridia : These are male sex organs present on the ventral surface of prothallus near
the rhizoids. Each antheridium is sessile, spherical structure and has a wall made up 3 cells.
The basal cell is called first ring cell; the middle cell is
 called is called second ring cell and the apical cell is called cap cell or lid cell. The
interior of the antheridium is filled up with few haploid androcytes (antherozoid
mother cells). At maturity, each androcyte metamorphoses inter spirally coiled,
haploid, multiflagellate antherozoids. The remaining cytoplasm turns
mucilaeinous.

Archegonia: These are borne on the central cushion-like area towards the notch on
ventral surface. Each archegonium is a sessile, inverted, flask-shaped structure
consisting of globular venter and tubular neck.
The venter is embedded in the prothallial tissue (cushion); hence it has no jacket of
its own. It contains single large egg or oosphere towards the base and venter canal
cell towards the neck. The neck is short and curved towards the posterior end (on
antheridial side). The wall of the neck is single layer in thickness with the cells
arranged in four longitudinal rows and closed at the top by lid cells. It contains a
single neck canal cell with two nuclei.
b) Fertilization : It takes place only in presence of water. As prothallus is protandrous
(antheridia matures first), cross-fertilization takes place. At maturity, in the
antheridium, protoplasts of androcytes metamorphose into antherozoids while the
remaining cytoplasm turns into mucilage. The mucilage absorbs water and swells
up to break open the lid cell to liberate antherozoids. The antherozoids swim in
water towards the archegonia of other plants.
At maturity, in the archegonium, neck canal cell and venter canal cell are dissolved
to form mucilage. It absorbs water; swells up and exert pressure on the lid cells to
break open the neck and to liberate the mucilage. The mucilage contains malic acid
which when gets dissolved in water, attracts
swarms of the
antherozoids towards
archegonia. This
phenomenon is called
chemotaxis. Many
antheozoids swim
through neck towards
the egg, but only one
fertilizes with it. As a
result of fertilization, a
diploid zygote is
produced, which
develops a protective
wall around it to form diploid oospore which eventually
develops into diploid sporophyte.
Water is essential for fertilization in fern because of
the following reasons:
i) When mucilage in the antheridium absorbs water, it swells up exerting pressure on
cap cell, which breaks open to liberate antherozoids. Thus for liberation of antheroids,
water is essential.
ii) Water is a medium for antherozoids to swim toward archegonia.
 iii) When mucilage in the archegonium absorbs water. It swells up to break open
lid cells, through which mucilage along with malic acid is liberated out. Thus,
for liberation of malic acid water is essential.
iv) When malic acid gets dissolved in water, it attracts swarms of antheroids
towards archegonia. The phenomenon is called chemotaxis for which water is
essential.
Young sporophyte

Many archegonia are fertilized but only one oospore develops into young
sporophyte. The oospore divides and redivides within the Venter to form embryo. The
early embryo shows foot, root, stem apex and cotyledon. The foot is muticellular and is
embedded in the gametophyte and absorbs nourishment for developing sporophyte.
Thus initially the sporophyte is dependant on gametophyte. Soon the root enters the soil
while the cotyledon and the stem apex bend, come up through the notch of the
prothallus and establish into young sporophyte. The prothallus gradually degenerate and
young sporphyte develops into fern plant.
Alternation of generations
In the iife-histories of
pteridophytes, there are two
. generations viz.; diploid asexual
generation called sporophyte and
haploid sexual generation called
gametophyte.

Sporophyte
produces gametophyte through
asexual reproduction and
gametophyte produces
sporophyte through sexual
reproduction. As these two
generation produce each other
alternately, i.e., the diploid
generation alternates with
haploid generation and so on,
hence, the process is called
alternation of generations.
In fern, sporophyte is
differentiated into roots, stem
and leaves. The roots absorb
 mineral salt and water while the
leaves prepare food material.
Thus, sporophyte is an
autotrophic, independent
generation.
Being asexual generation,
it produces sori on abaxial side of
leaflets. Each sorus contains group of sporangia, which produces haploid spores after
meiosis. The spores germinate into gametophyte.
 The gametophyte called prothallus is a heart-shaped structure with 5mm diameter. It
shows presence of chlorenchymatous cells to prepare food material and rhizoids to absorb
mineral salts and water. Thus, in spite of its smallness, prothallus is also an autotrophic,
independent generation.
Being sexual generation, it produces sex organs, which are produced on its ventral
surface. The male sex organs called antheridia produces male gametes called antherozoids
whereas the female sex organ called archegonia produces female gametes called eggs. Male
gamete fuses with the female gamete to form diploid zygote, which eventually develops into
diploid sporophyte.
Thus, the diploid sporophyte produces haploid gametophyte and the haploid
gametophyte produces diploid sporophyte. Hence, diploid phase alternates with haploid
phase and the process is called alternation of generations.

~STELAR EVOLUTION ~
The word 'Stele' is derived from a Greek word meaning 'Column or Strand' and is
applicable only to the primary vasculature. In 1886, Van Tieghem and Douliot proposed
that the fundamental parts of a shoot are a cortex and a central core of vascular cylinder or
stele and that the two delimited from each other by the endodermis. Therefore, stele is
defined as "the core of axis which includes the vascular system, pith, interfascicular portion
and pericycle. Regarding the position of endodermis there are two views. According to one
view, it is the innermost layer of cortex, whereas, according to the other view, it is the
outermost layer of the stele. Four basic evolutionary categories of steles are recognized,
viz.; Most primitive types of steles (Protosele), Advanced types of steles (Siphonostele),
More advanced types of steles (Solanostele) and Most advanced types of steles
(Atactostele). A brief account of their evolution is given below : 1. Most Primitive Types
of Steles
Protostele : As the name suggests, it is the most primitive type of stele consisting of a
solid cylinder of xylem in the centre, surrounded by phloem which is in turn surrounded by
single layered pericycle. Pith is absent, e.g., fossil plant Rhynia (aerial branch) and some
living plants like Psilotum (rhizome), Nephrolepis (rhizome), etc. It is further divided into
following types :
a) Haplostele : It is considered as most primitive type of protostele. The core of xylem
is smooth. It appears circular in a cross section and is surrounded by phloem. The
origin of the xylem is exarch as protoxylem is situated at the periphery of the xylem,
e.g. stem of Lycopdium and Selaginella.
b) Actionstele : In this, the pith is absent (protostele) and the centre is occupied by
xylem. Hence, it is a protostele. However, the xylem-core has radiating arms
forming a stellate or star-shaped structure (actinos - star). The xylem is exarch in
origin as the protoxylems are situated at the end of the radiating arms. Phloem
completely surrounds the xylem and is in turn surrounded by pericycle and
endodermis, e.g., stolon of fern, stem of Lycopodium sermtum.

c) Plectostele : The xylem and the phloem appear intermingled. The radiation of
arms of the xylem splits to such an extent that they appear as separate parallel
plates. The phloem completely surrounds xylem and is in turn surrounded by
pericycle. Such protostele having plate-like, exarch xylem is called plectostele,
e.g., stem of Lycopdium annotinum and Lycopdium clavatum.
d) Mixed stele : It is considered as the most advanced type of protostele which consists of
irregular groups of xylem surrounded by phloem.
It appears as if xylem is embedded in the phloem.
The vascular tissues are surrounded by pericycle
and endodermis. Such stele is called mixed stele, e.g.,
stem of Lycopdium cernuum.
2. Advanced Types of Steles
Siphonostele ; In this type, pith is present at the
centre of the stele in the form of a cylinder.
Parenchymatous pith is surrounded by xylem,
phloem, pericycle and endodermis. The presence
of the pith is an evolutionary character which
changes a protostele to a siphonostele.
Siphonostele is further divided into following
types :
a) Ectophloic siphonostele : In this type of stele, there is pericycle and endodermis
only on the outer side of the vascular tissues, i.e., xylem and phloem and phloem
is only on the outer side (ecto) of the xylem. The centre is occupied by pith, e.g.,
stem of Osmunda and Schizaea.

b) Amphiphloic siphonostele : In this, xylem forms a ring. Phloem is present on

both the sides of the xylem ring and is known as outer phloem and inner phloem.
Outside the outer phloem, there is outer endodermis and outer pericycle and
inside the inner phloem, there is inner pericycle and inner endodermis. Xylem
ring is formed of metaxylem while there are several patches of protoxylem on the
outer face of the xylem ring (exarch). Thus, there are two pericycles, two
endodermes, two groups of phloem and a single group of xylem. The pith is
present in the centre, e.g., Rhizome of Marsilea.
The above two types of stele exhibit a continuous cylinder of vascular tissue and is not
as evolved as the subsequent type which has a network of vascular bundles.
3. More Advanced Types of Steles
Solenostele : This type is more advanced than the former two cited above. When a
siphonostele is interrupted by leaf gaps, it is termed as a solenostele. It is further divided
into following two types:
a) Dictyostele : In it, the siphonostele gets dissected by several overlapping leaf gaps
and results in the formation of intervening areas of vascular tissue, called
Meristeles. Each meristele consists of xylem, phloem, pericycle and endodermis,
e.g., Rachis of Nephrolepis (fern).

b) Eustele : It is the modification of the siphonostele, in which, the vascular cylinder

is dissected and consists of collateral or bicollateral strands, with leaf gaps and
interfasicular areas. The leaf gap and inter-fascicular region cannot be delimited.
The dissected parts of the stele are called vascular bundles. These vascular
bundles are arranged in the form of ring. There is endodermis which is common
. to all the vascular bundles and is present on the outer side. Pith is present at the
centre. This type of a stele occurs in highly evolved vascular plants like
gymnosperms and angiosperms (dicotyledonous plants), e.g., Sunflower stem.

4. Most or Highly Advanced Type of Stele

Atactostele : It is the most complex and highly evolved stele, in which, the vascular
cylinder consists of widely scattered vascular bundles in the ground tissue. These vascular
bundles are not delimited by endodermis and pericycle as both are absent. Such stele
having a widely scattered network of vascular bundles is called atactostele (Atactos
meaning without any order). It is the characteristic of monocotyledonous plants, e.g.,Maize
stem.