PTERIDOPHYTA

AIPMT Syllabus: General Characters, Eusporangiate and Leptosporangiate type of development,

Classification, Stelar system in Pteridophytes, Life cycle of Selaginella, Life cycle of pteris.
IPE/EAMCET Syllabus: Life cycle of pteris.

PTERIDOPHYTA
 The pteridophytes (Gk. pteron = feather, phyton = plant, Haeckel, 1866) are the spore bearing
most primitive vascular plants. They are commonly called vascular cryptogams.
 Oswald Tippo (1942) has placed them in tracheophyta. The pteridophytes are an assemblage
of flowerless, seedless, spore bearing plants that have successfully invaded the land. They
were perhaps the first land plants to evolve during ordovician (425-450 million years back)
age.
 Today the group is represented by about 11,000 living species.

THE SALIENT FEATURES OF PTERIDOPHYTES

Occurrence
 The pteridophytes grow under varied habitats, most of them are terrestrial plants and thrive
well under damp and shady conditions while some flourish well in open grasslands or even
under xeric conditions (e.g., Selaginella lepidophylla). A few pteridophytes grow as aquatic or
semi aquatic (e.g., Marsilea. Azolla and Salvinia) while still others are epiphytes (e.g.,
Lycopodium phlegmaria).

The Plant Body (The Adult Sporophyte)

 The main independent plant body of pteridophytes is sporophyte (2n).
 The sporophytic plant body is cormophyte differentiated into true root, stem and leaves. Some
primitive members of the group may lack true root and well developed leaves e.g., order
Psilophytales and Psilotales. In these, roots are replaced by rhizoids and stem axis becomes
photosynthetic.

Fig 1. Selaginella kraussiana – A. general habit, B. A part of the plant,

C. Small portion of B showing arrangement of leaves.

 The primary root is short lived. It is replaced by adventitious roots which generally arise
endogenously.
 Plants generally exhibit dorsi ventral or radial symmetry. The .branching of stem may be
dichotomous or monopodial.
 The stem bears leaves which may be small microphyllous (e.g., Lycopodium, Equisetum) or
very large macrophyllous (Pteridium, Pteris and other ferns).
 All the vegetative parts possess vascular tissues, organized in definite groups or stele. The
stele, in pteriodophytes may be simple protostele (e.g., Lycopodium, Selagillella), medullated
protostele or siphonostele (e.g., Marsilea) or dictyostele (Many ferns) or even eusteles (e.g.,
Equisetum)

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 The Xylem mainly consists of tracheids and xylem parenchyma. The vessels are generally
absent but primitive types of vessels have been reported in Selaginella, Equisetum,
Pteridium, etc.
 The phloem consists of sieve-tube and phloem parenchyma. The companion cells are absent.
 Secondary growth does not occur in most of the living pteridophytes Isoetes being an
exception

REPRODUCTION
 The sporophytic plant reproduces by means of spores that are produced in small capsules
called sporangia.
 Leaves bearing-the sporangia are called sporophylls.
 The sporophylls may be widely scattered on a plant or may be clustered in definite areas and
structures called strobili (e.g., Selaginella and Equisetum).
 In aquatic pteridophytes, the sporangia are produced within the specialized structures called
the sporocarps (e.g., Marsiliea, Salvinia and Azolla).
 According to the mode of development, the sporangia are of two fundamental types, the
eusporangium and leptosporangium. The eusporangium is found in most cases and develops
from several sporangial initials (e.g., Psilotum, Lycopodium, Selaginella, Equisetum, etc). The
leptosporangium is found only in some of the advanced ferns and develops from a single
superficial cell (e.g., Marsilea, Pteridium, Pteris, etc.).
 Within the sporangia are developed the diploid spore mother cells or sporocytes. These spore
mother cells undergo meiosis or reduction division to form spores. If all the spores are of the
same size, the plant is said to be homosporous (e.g., most of pteridophytes) and if they are of
two different sizes, the plant is called heterosporous (e.g., Selaginella, Isoetes, Marsilea,
Salvinia, Pilularia, Regnelidium, Styiltes and Azolla). In the heterosporous types, the two
different types of spores are produced in separate sporangia. The smaller spores are termed
as microspores and are developed in microsporangia, while the larger spores that are
generally produced in smaller numbers, are termed as megaspores and are formed in
megasporangia.

THE GAMETOPHYTE
 The spores, on germination give rise to the haploid gametophytes or prothalli that are usually
small and insignificant structures. The gametophytes are inconspicuous as compared to the
sporophytes. One of the most characteristic features of the pteridophytes is that the
sporophyte has become the dominant morphological part of the life cycle while the
gametophyte has been much reduced.
 The gametophytes are of two general types. Gametophytes that develop from homospores
and grow upon the soil (outside the spore wall) to form independent plants, are known as
exosporic gametophytes (e.g., Psilotum, Lycopodium sp. and Ophioglossum). Gametophytes,
that develop from heterospores and most of the part retained within the original spore case,
are called endosporic gametophytes (e.g., Selaginella, Isoetes and Marsilea).
 Exosporic gametophyte is typically a delicate, thin thallus and is commonly called the
prothallus. In most of the vascular cryptogams, the exosporic gametophytes grow exposed to
light and remain attached to the ground by numerous rhizoids. In such cases they
manufacture the food by means of their chloroplasts and live an independent life. The rhizoids
are meant for the absorption and fixation.
 In some vascular cryptogams, the exosporic gametophytes are devoid of chlorophyll and are
subterranean in habit. In such cases they obtain their food by symbiosis through the agency
of mycorrhiza that occurs within the tissue of the prothallus or gametophyte, e.g., Psilotum,
some species of Lycopodium and Ophioglossum.
 Endosporic gametophytes that develop from heterospores are greatly reduced structures.
They develop largely or entirely within the spore wall and live on food deposited in the spores.

SEX ORGANS
 The gametophyte or prothallus bears the sex organs, the antheridia and archegonia.
Typically, the gametophytes formed from the homospores are monoecious, i.e., both
antheridia and archegonia are borne in large numbers on the same gametophyte or
prothallus. The gametophytes formed from the heterospores are dioecious, the antheridia and
archegonia developing on separate male and female gametophytes.

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Antheridia: The antheridia may be embedded either wholly or partially in the tissue of
gametophyte or they are projected from it. At maturity, each antheridium is a globular
structure. It consists of an outer single cell layered sterile wall inside which are found a large
number of androcytes. Each androcyte gives rise to a single motile antherozoid.
Archegonia: The archegonia in pteridophytes resemble closely with those of the bryophytes.
Each archegona is a flask shaped structure, consisting of a basal swollen, embedded portion
the venter and a short neck. The neck consists of 4 vertical rows of cells, each with 2 to 4
cells. The neck has a single binucleate neck canal cell (14 neck canal cells in Lycopodium).
The venter encloses the egg and ventral canal cell. At maturity, the apical cells separate, the
neck canal cell and venter canal cell disintegrate forming a passage for the antherozoids to
reach the egg cell.
Fertilization: Fertilization in all cases is accomplished by the agency of water. The mucilage
and malic acid, formed by the disintegration of neck canal cell and venter canal cell, not only
provide a medium for swimming of antherozoids but also chemotract them. Many
antherozoids enter into the venter but only one, the most active one, fuses with female
gamete. The fusion of a male gamete and an egg restores the diploid chromosome (2n)
number and results in the formation of the zygote.

THE EMBRYO (THE NEW SPOROPHYTE)

 The zygote undergoes repeated divisions to form a new sporophyte. The young sporophyte
remains attached to the gametophyte by means of a foot and draws nourishment from the
prothallus (gametophyte) until it develops its own stem, roots and leaves. The sporophyte is
dependent on the gametophyte only during its early stages.

STELAR SYSTEM IN PTERIDOPHYTES

 At one time the vascular bundle was considered the fundamental unit in the vascular system
of the pteridophytes. Later a stelar theory was proposed by Van Tiegham and Douliot in 1886.
According to this theory the primary structure of the stem and root are basically similar and
the fundamental parts of each are the cortex and a central column, the stele, and that the two
are separated from each other by the endodermis. According to the proponents of this theory,
the endodermis constitutes the innermost layer of the cortex, and pericycle the outermost
portion of the stele. The stele includes not only the primary vascular tissues but also the
pericycle and pith (if present). The idea of the stelar theory was widely accepted by many
scientists and proved to be useful in the comparative anatomical and phylogenetic studies of
the vascular plants.

DIFFERENT TYPES OF STELE FOUND IN THE PTERIDOPHYTES

 Jeffery (1898) viewed the stellar organization from the point of view of evolution.
 Two main types of steles are found in the pteridophytes, protostele and siphonostele.

PROTOSTELE (Jeffery, 1903)

 It is the simplest and most primitive type of stele in which the vascular tissues form a central
solid mass of xylem surrounded by phloem, pith being absent. Such a stele has been
regarded as the most primitive from a phylogenetic stand point from which other types of
steles would have evolved in the course of evoulutionary specializatioon. The protostele has
been classified into the following two types:
 Haplostele (Brebner, 1902): A protostele with central solid and smooth core of
xylem surrounded by phloem is known as a haplostele. This particular type of
protostele has been regarded as the most promititve among the different types. It
occurred in primitive psilo like Rhynia and Horneophyton and is found in a number of
living genera, e.g., Lycopodium cernuum and Selaginella craussiana.
 Actinostele (Brebner, 1902): In a number of pteridophytes, the central xylem core of
a protostele is not smooth but is thrown into radiating ribs with phloem alternating with
its rays when seen in a cross section. Such a type of protostele is termed an
actinostele. It is .found in Psilotum triquetrum and Lycopodium serratum.
 In the stems of some species of Lycopodium, e.g., L. clavatum and L. volubile, when
seen in a cross section the xylem occurs in the form of small parallel bands
alternating with the phloem plates. This specialized form of actinostele is usually
telmed asplectostele (Zimmerman, 1930).

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 In Lycopodium cernuum, the xylem when seen in a cross section appears in the form of
irregular groups that are embedded in the ground mass of phloem. This type of actinostele is
called the mixed protostele. In Hymenophyllum demissum, mixed protostele with pith is found.

SIPHONOSTELE (Jeffery, 1893)

 A kind of stele in which pith is present in the central region is called a siphonostele or
medullated protostele. This type of stele is said to have been evolved from the protostele, and
represent an evolutionary advance stage. In siphonostele, the vascular tissues are arranged
in the form of a hollow cylinder, with distinct pith in the centre. It is found in the stems of most
members of the filicophyta.
 Transition of protostele into siphonostele is seen in plants like Botrychium, Gleichenia,
Osmunda, Schizaea, etc.
 Types of siphonostele: According to the distribution of the vasculartissues, the siphonostele
has been classified into following types:
 Ectophloic siphonostele
 Amphiphloic siphonostele
 Cladosiphonic siphonostele
 Solenostele
 Dictyostele or dissected siphonostele.
 Polycyclic siphonostele
 In the ectophloic siphonostele, the xylem is in the form of a hollow cylinder surrounding pith
with the phloem occurring only outside the xylem. It is found in Equisetum and some ferns like
Osmunda and Schizaea.
 In the amphiphloic siphonostele the xylem forms a hollow cylinder enclosing the pith with the
phloem occurring both on the inner side and outer side of the xylem. It is found in ‘the ferns
like Adiantum and Marsilea.
 In the lower vascular plants, e.g., Lycopodium and Selaginella the leaf gaps are absent. In
these plants the stele shows a continuous ring of tissues. A siphonostele that has no leaf gap
is termed as cladosiphonic siphonostele. In ferns large leaf gaps are found in the
siphonosteles. Such a siphonostele with the leaf gaps is known as phyllosiphonic
siphonostele.
 In some ferns there may be leaf gaps that do not overlap. A cross section through the
internode shows a continuous ring of vascular tissues. This particular type of siphonostele in
which there is no overlapping of gaps is termed solenostele. It may be ectophloic solenostele
or amphiphloic solenostele.
 In some ferns, the leaf gaps are large and extend vertically so that they do overlap. As a
result of overlapping of the leaf gaps the vascular tissue of the stem appear as scattered
series of vascular bundles. This type of siphonostele is known as dictyostele or dissected
siphonostele (Brebner, 1902). In this type the vascular strands are interconnected and form a
cylindrical meshwork. When seen in a cross section this type of stele shows the presence of
separate strands called meristeles. Each such strand or meristele is concentric in structure
consisting of a central strip of xylem surrounded by phloem. Dictyostele is found in Pteris,
Pteridium, Polypodium and Dryopteris
 In some pteridophytes, the gaps occur in the steles that are not associated with the leaf
traces. Such gaps have been termed perforations. Therefore a dictyostele having such gaps
or perforations is termed as perforated dictyostele.
 In certain pteridophytes, complex type of stelar structure is seen in which two or more
concentric rings of vascular tissues are present. This type of stele is known as polycyclic
stele. In Pteridium aquilinum, there are two concentric rings of vascular tissues. in which the
inner ring forms a siphonostele and the outer ring forms a dictyostele consisting of numerous
meristeles. In Matonia pectinata there are three rings of vascular tissues.

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Fig 2. Diagrammatic representation of steles. A. Haplostele; B. Actinostele; C. Plectostele; D.

Mixed protostele with phloem; E. Mixed protostele with parenchyma; F. Ectophloic
siphonostele; G. Amphiphloic siphonostele; H. Ectophloic solenostele; I. Amphiphloic
solenostele; J. Dictyostele K. Polycyclic siphonostele; L. Polystele.

LIFE CYCLE OF PTERIDOPHYTES

 The life cycle of a pteridophyte comprises of two distinct phases or generations, each of
which produces the other. One phase or generation, the sporophyte, is diploid (2n) and the
dominant part of the life cycle. It is the part of the life cycle in which vascular tissue is
developed. The sporophyte develops sporangia within which are diploid cells called spore
mother cells or sporocyte. Each spore mother cell divides by meiotic division. As a result of
meiosis, spores are produced which are, therefore, haploid (n).
 The gametophytic generation is always small, inconspicuous and bears male and female
gametes. The male gamete i.e. antherozoids are produced in large number within the
antheridium. The female gamete i.e. egg is generally borne single within the archegonium.
Fertilization takes place in the presence of water when an antherozoid fuses with an egg to
produce a diploid zygoyte. The zygote germinates to form the new sporphyte. This generation
is called sporophytic generation.
 Thus, the life cycle of a pteridophyte consists of an alternate succession of sporophytic and
gametophytic generations (heteromorphic or heterologous alternation of generation).

Heterospory and Seed Habit in Pteridophytes

 The occurrence of two kinds of spores in the same plant is called heterospory. Of the two
kinds of spores, the smaller ones are called microspores and the larger ones are termed as
megaspores and are produced within the microsporangia and megasporangia, respectively.
 The production of two kinds of spores in heterosporous plants is definitely related with a
differentiation in sex since a microspore develops,a male gametophyte and a megaspore a
female gametophyte. The heterospory, thus, is associated with the sexual differentiation of
gametophyte.

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 The differentiation of spores into microspores and megaspores and their dependence on the
parent sporophyte for the nutrition are the certain features in the life cycle of Selaginella that
have been considered as the essential prerequisites for the formation of seeds, characteristic
of spermatophytes. It is generally agreed that the seed plants arose from the heterosporous
vascular plants that instead of discharging the megaspore acquired the habit of retaining it
within the megasporangium.
 In the seed-bearing plants there are two kinds of spores: microspores and megaspores that
grow to form male and female gametophytes, respectively; In these plants, the single
megaspore is not shed from the megasporangium but is retained within it while still attached
to the mother plant. It germinates inside the megasporangium (nucellus) producing the much
reduced female gametophyte bearing the archegonia. Later the nucellus and the
gametophyte are protected by a covering or integument and the whole structure is known as
an ovule. The female gametophyte gets nutrition for its development from the parent plant
and thus does not need to produce by its own effort. After fertilization the zygote within the
ovule gives rise to an embryo, the rest of the gametophytic tissue gives rise to the nutritive
tissue or endosperm and the integument thickens to form a seed coat. This entire structure,
i.e., the integumented ovule is known as seed. It is detached from the parent plant and
germinates to form a new plant.
 Thus we find that for the production of seeds, the following prerequisites are essential:
 Production of two types of spores (heterospores).
 Megasproangium (nucellus) does not open and the single megaspore within it
germinates to from the female gametophyte.
 Nucellus becomes invested by a covering or integument which later forms a seed
coat.
 Within the nucellus is formed a linear tetrad of four haploid megaspores as a result of
the reduction division in the functional megaspore mother cell. Out of these four
megaspores, the lowermost gives rise to the female gametophyte whereas the rest
degenerate.
 The male gametes reach the egg by means of a tubular outgrowth of the male
gametophyte known as the pollen tube.
 Fertilization and formation of embryo take place within the megasporangium.
 The embryo undergoes a resting period.
 Selaginella exhibits a remarkable approach to the seed habit the characteristic of the
spermatophytes because of the following features:
 The heterospory occurs in almost all the species of Selaginella.
 In most species only one functional megaspore mother cell is produced which by
reduction division.
 One megaspore is formed in each megasporangium and this single megaspore is not
shed but germinates to form the female gametophyte.
 The fertilization and development of embryo in both the species take place while the
megaspore is enclosed within the megasporangium.
 Therefore, it becomes evident that Selaginella is considerably advanced towards the seed
habit in few species but its approach to the true seed is not completed due to the following
features:
 The megasporangum lacks an integument or covering.
 The retention of the megaspore permanently within the megasporangium has not
become evident.
 After the development of the embryo there is a lack of resting period.
 Economic importance:
 It is a good source of potash when burnt.
 Ferns are grown as ornamental plants because of their graceful foliage.
 An antihelmintic drug is obtained from rhizome and petiole of Dryopteris.
 The sporocarps of Marsilea are used as food by certain tribes.

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Classification of Peteridophyta
 The Pteridophyta is divided into four classes, viz. Psilopsida, Lycopsida, Sphenopsida and
Pteropsida, on the basis of organization of plant body including the nature of leaf, vascular
system, and location of sporangia.

PTERIDOPHYTA

Psilopsida Lycopsida Sphenopsida Pteropsida

Distinguishing characters of these sub-phyla are described below.

I. Sub-Phylum – Psilopsida
(1) These are the oldest known vascular plants, most of them (except Psilotum and
Tmesipteris) are fossils.
(2) Plant body is relatively less differentiated.
(3) Roots are absent, instead dichotomously branched rhizome is present.
(4) Aerial axis is either naked or have small spirally arranged leaves.
(5) Sporangia are cauline (i.e., directly borne on the axis or stem); they are lateral or
terminal in Position.
Examples. Psilotum,Tmesipteris
II. Sub-Phylum – Lycopsida
(1) Plant body is differentiated into root, stem and leaves.
(2) Leaves are small (i.e., microphyllous) with a single unbranched vein.
(3) Sporangia develop in the axil of the Sporophylls
(4) Sporophylls generally form compact strobili. Examples. Lycopodium, Selaginella etc.
III. Sub-Phylum — Splicuopsida
(1) Stem differentiated into nodes and internodes.
(2) Leaves microphyllous, present in whorls at each node.
(3) Sporangia are borne on the sporangiophores which form compact cones at the apex
of the fertile branches.
Example: Equisetum
IV. Sub-Phylum — Pteropsida
(1) Plant body well differentiated into root, stem and leaves.
(2) Leaves megaphyllous, Pinnately compound.
(3) Sporangia develop on the ventral surface of the Sporophylls, usually aggregated into
sori.
Examples : Dryopteris Pteris, Pteridium, Polypodium etc.

DRYOPTERIS: LIFE CYCLE

 There are 150 species of Dryopteris; about 25 species have been reported in India. The most
common species is Dryopteris fluxmas, which is commonly known as Beech fern or male
shield fern or Hay scented fern. It is found in moist and shady places in tropical, sub-tropical
and temperate regions.
 The plant body is a sporophyte, which is differentiated into root, stem and leaves. The roots
are adventitious. The stem is dark brown underground rhizome growing obliquely. From upper
surface of rhizome many leaves arise acropetally. The leaves are large, called fronds, and are
bipinnately compound. The young leaves show circinate vernation. Persistent leaf bases of
the dead leaves are present in the old part of rhizome. The younger part of rhizome and
rachis are covered with multicellular brownish scales called ramenta.
 Reproduction takes place by vegetative methods and by spores.

VEGETATIVE REPRODUCTION
 Vegetative reproduction takes place by adventitious buds that develop on the rhizome. These
buds give rise to new plants. Besides this, fragmentation of rhizome also helps in vegetative
propagation.

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SEXUAL REPRODUCTION
SPORE PRODUCING ORGANS
 Spores are formed within sporangia. They develop on the ventral (lower) surface of ordinary
foliage leaves. The leaves bearing sporangia are known as sporophylls. Many sporangia arise
irregularly from placental tissue developed at the tip of an ultimate vein. The sporangia of all
developmental stages are aggregated in cluster called sori. These are arranged in two rows,
one on each side of the main’ vein. Thus sori are sub marginal and discontinuous. Sorus is
covered by a thin membranous shield like or kidney shaped outgrowth of the leaf, called
indusium

Fig 3. Dryopteris. A. Part of a sporophyll with sori, B. T.S. of sorus.

 Each sporangium develops from a single initial cell. The development is of leptosporangiate
type. Mature sporangium consists of a stalk and a body or capsule. The stalk is long, slender
and multicellular. The capsule is lens shaped. The capsule wall, is only one cell in thickness.
A row of thin cells on one side of the capsule marks the line of dehiscence. It is called
stomium. Besides this, a ring of modified cells, called annulus extends along the edge of the
capsule. The cells of the annulus have thick radial and inner walls while the outer tangential
walls are thin.
 Capsule encloses 8 or sometime 16 spore mother cells. Each of them divides meiotically and
forms haploid spores. A mature sporangium so has 32 or 64 spores. The spores are the first
cells of the new gametophytic generation. All the spores are similar in shape and structure,
So, Dryopteris is a homosporous fern. The spores are asexual reproductive bodies.
 The spore wall is two layered: an outer thick ornamented exine and inner thin and smooth
intine. Each spore has a single haploid nucleus.
 Dispersal of spores: Dispersal of spores takes place when the atmosphere is dry. In such
conditions the indusium dries and shrivels and the sporangia are exposed to the dry
atmosphere. The annulus and stomium help in the dehiscence of sporangium by a purely
mechanical action. In dry atmosphere water evaporates from thin outer wall of the annulus
which are pulled in and the thick radial walls contractThis causes the capsule to break open
transversely at the stomium. In this process the upper half of the sporangial wall swings
backward along with annulus in the form of a cup. At this stage almost all the spores are
lodged in this cup. Later, due to continued drying, the annulus snaps back to its original
position. This tosses the spores out into the air.

GAMETOPHYTE
THE PROTHALLUS
 The spore is the mother cell of gametophytic generation It germinates when temperature and
moisture are suitable. As a result of moisture absorption, exine ruptures and intine comes out
in the form of a germ tube. It develops chloroplasts and divides transversely to form a green
filamentous structure resembling moss protonema. The germ tube attaches itself to the soil by
rhizoids. At a very early stage in development, the uppermost cell of the filament divides and
establishes an apical cell. It cuts off cells alternately on two sides till a heart shaped
gametophyte is formed. Mature gametophyte is thin flat and green structure, approximately
1/4 inch in diameter. It is known as prothallus. The apical part of prothallus has an apical
notch where the growing point is situated. The lobes of prothallus are only one cell in
thickness whereas the central part lying immediately below the apical notch is several celled
thick. Many one-celled rhizoids develop on the ventral surface of the thallus and serve as
organs of attachment.

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Fig 4. Detailed structure of a prothallus showing position of archegonia, antheridia and

rhizoids

 All the cells of the prothallus, except rhizoids are green and so, the gametophyte is
autotrophic.

SEX ORGANS
 The male and female reproductive structures are antheridia and archegonia, respectively. The
prothalli are monoecious, homothallic and protandrous (antheridia maturing earlier than
archegonia). Both the sex organs are produced on the ventral surface of the thallus.

ANTHERIDIA
 The antheridia develop in the basal region of the prothallus, among the rhizoidal cushion.
Matured antheridium is a dome shaped structure which is projected beyond the surface of the
prothallus. The wall of the antheridium is composed of only three cells. Two cells form a ring
around antheridium and are known as first ring cell and second ring cell. The third cell, which
forms a cap of the antheridium, is known as cap cell. In each antheridium, there are usually
32 spirally coiled multiflagellate antherozoids. The antherozoids of ferns thus resemble with
those of Cycas in their multiflagellate nature.

ARCHEGONIA
 The archegonia are produced in the thickened portion of the prothallus,just behind the apical
notch. Mature archegonium is a flask shaped structure. It has a basal enlarged venter that is
deeply sunken in the tissue of the prothallus and a neck that project beyond the surface of the
prothallus. The wall of the neck consists of four rows of neck cells. The venter has a large egg
and a small venter canal cell. The neck canal of the young archegonium is occupied ‘by an
axial formation between the two neck canal cells and, therefore, almost invariably there is a
single binucleate neck canal cell.

FERTILIZATION
 Fertilization takes place when the antheridium absorbs water. This creates turgor pressure
that pushes the cover cell of the antheridium and the antherozoids are set free. Almost at the
same time the venter canal cell and the neck canal cell of the archegonium disintegrate to
form a mucilaginous substance. It contains malic acid that attracts antherozoids towards the
neck of the archegonium. This is a chemotactic movement. Many antherozoids swim into the
neck of the archegonium, but only one of them fuses with the egg. The fusion results in the
formation of a diploid zygote or oospore. The new sporophytic phase begins with the zygote.

EMBRYO AND THE YOUNG SPOROPHYTE

 Generally, many archegonia are fertilized but only one of them develops into embryo. So, only
a single sporophyte is found attached to each prothallus. Soon after fertilization, the zygote
secretes a thick wall and begins to grow. The first division of zygote is vertical, i.e., along the
long axis of the archegonium. The resulting two hemispherical cells divide till eight cells are
formed. These are arranged in two tiers of four cells each. It is called octant stage. The upper
tier, known as epibasal tier, forms primary leaf (or first leaf) and primary root. The lower of the
hypobasal tier forms foot and the embryonal stem. Subsequent divisions in the embryo are
irregular. The foot, which is the first organ to develop, is a short, projecting spherical mass of
cells. It absorbs food, water and minerals. The young sporophyte is, totally dependent upon

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gametophyte in the beginning. The primary leaf, primary root and embryonal stem develop
later.
 After the establishment of foot, the primary leaf and primary root, the embryo emerges from
the archegonial wall. The embryonal stem forms rhizome that gives out a number of
adventitious roots. The primary leaf is replaced by a bipinnately compound leaf. Thus, a new
independent sporophyte is established and the prothallus dries up and disintegrates. The
young sporophyte now manufactures its own carbohydrate food and absorbs water and
minerals from the soil.

Fig 5. Diagrammatic life cycle of Dryopteris

ALTERNATION OF GENERATION
 Dryopteris has a heteromorphic alternation of generation. There are two morphologically
distinct phases in its life cycle: the sporophytic phase and the gametophytic phase, which
alternate with each other. The sporophytic phase that forms spores is long lived and is
differentiated into roots, stem and leaves. The gametophyte, on the other hand is relatively
short lived and thalloid. It manufactures its organic food by itself. The sporophyte is
dependent upon gametophyte when young but becomes independent as it attains maturity.
 The sporophyte is diploid. The diploid spore mother cells undergo meiosis and form haploid
spores. The spore is the mother cell of the haploid gametophytic generation and forms
gametophyte which produces haploid gametes. Haploid male and female gametes fuse to
form diploid zygote, which is the mother cell of the sporophytic generation. The zygote grows
into embryo and develops into sporophyte.

SELAGINELLA : LIFE CYCLE

 Selaginella, commonly known as spike moss or little club moss prefers moist, cool and shady
places, though a few species (e.g., S. lepidophylla and S. rupestris) are xerophytes.
The .xerophytic species roll up to form ball-like structures (cespitose habit) during dry season
but spread out and revive when sufficient moisture is available (e.g., S. lepidophylla). Such
plants are called resurrection plants. S. bryopteris is sold in the market under the name
“Sanjivani”.

EXTERNAL FEATURES
 The main plant body represents the sporophyte (2n), and is an evergreen herb differentiated
into root, stem and leaves.
 The stem is erect prostrate and dichotomously branched. Positively geotropic structures
called rhizophores arise from the stem at the point of branching. Rhizophore bears
adventitious roots at its tip. Rhizophores are non-green, leafless thread like structures. They
lack root hair and root cap. The rhizophores develop exogenously from meristem (a special

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meristem) which lies at the point of branching. Since rhizophore resembles the stem in some
characters, and the root in other characters, it was regarded as organ suigeneris (organ of
independent origin) by Bower and Goebel.
 The leaves are small, simple, sessile called microphylls (being single-veined).
 The leaves are ligulate (i.e., bear a small multicellular scale-like structure) at the base of the
leaf on the adaxial side). The bulbous basal region of the ligule is made up of larger cells
called glossopodium.

Fig 6. Sporophyte of S. kraussiana

 On the basis of types of leaves present, the genus Selaginella consists of two sub-genera:
 Sub-genus Homoeophyllum: All leaves are of same type (e.g., S. rupestris); stem is
erect.
 Sub-genus Heterophyllum: Leaves of two sizes, different sizes (heterophylly);
arranged in four vertical rows on the stem, e.g., S. kraussiana.

INTERNAL ORGANISATION
 Stem: Anatomically stem has an outer parenchymatous epidermis, a few layers of
sclerenchymatous hypodermis, parçnchymatous cortex and one or more steles. Thus the
stem is polystelic with 1 to 16 steles depending upon the species (S. spinulosa - 1 stele; S.
kraussiana - 2 steles; S. laevigata - polystelic). Each stele is a protostele and is surrounded
by air space.
 The stele is connected to the cortex across the air space by radially elongated filament-like
trabeculae. The trabeculae show casparian strips; hence, are believed to be endodermal
cells. The xylem is diarch, exarch and made up of tracheids, though a few vessels are present
in S. rupestris.
 Root: Root has a layer of epiblema with occasional root hair. It is followed by cortex,
endodermis, pericycle and a single monarch exarch xylem surrounded by C-shaped
phloem.
 Rhizophore: Rhizophore has an outer thick walled epidermis without root hair,
sclerenchymatous hypodermis, thin walled cortex, endodermis, pericycle and phloem
surrounding mesarch xylem on all sides.
 Leaf: Leaf has an upper and lower epidermis with stomata. Mesophyll is not
differentiated. Its cells contain one or more cup-shaped chloroplasts having granoids
(having irregularly stacked thylakoids). One vascular strand with protostelic condition
occurs in the mid-rib region. It is covered by bundle sheath.

REPRODUCTION
 Selaginella reproduces vegetatively and sexually.

VEGETATIVE REPRODUCTION
 Vegetative reproduction occurs by fragmentation of bulbils (e.g. S. subdiaphana) and stem
tubers (S. chrysocaulos)

SEXUAL REPRODUCTION
 It takes place by spores. Being heterosporous, Selaaginella produce two types of spores
microspores and megaspores produced-in microsporangia (producing numerous micro-
spores) and megasporangia (producing usually only four megaspores).
 The microsporangia and megasporangia are borne on microsporophylls and
megasporophylls, respectively, both of which aggregate to form cone (or strobilus) at the
apices of branches.

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 The sporangia are eusporangiate (developing from a group of initial cells).

 Each sporophyll is ligulate and bears only one sporangium at its base on the adaxial surface.
 Microsporangium consists of a multicellular stalk and a round/oval body covered by two-
layered jacket. A single-layered tapetum present beneath the jacket, provides nourishment for
developing spores. Many diploid microspore mother cells are formed in each sporangium
which undergo meiosis to form numerous microspores.
 Megasporangium is large four-lobed structure (one megaspore per lobe) having two-layered
jacket and a multicellular stalk. In S. monospora, only one megaspore survives. Megaspores
are formed by meiosis in megaspore mother cell.

GAMETOPHYTIC GENERATION
 Microspore develops into male gametophyte which is highly reduced. In initial stage it is 13-
celled consisting of one prothallial cell plus 12 cells of the antheridium (8jacket cells + 4
primary androgonial cells.). The microspores are liberated at 13-celled stage. The primary
androgonial cells givide (repeatedly to form 128 or 256 androcytes, each metamorphoses into
an oval biflagellate’ antherozoid.
 The megaspore germinates in situ (even before dispersal) to form an autotrophic multicellular
female gametophyte (female prothallus).
 After the release of megaspores at this multicellular stage, a few superficial cells act as
archegonial initials each of which develops into an archogonium.
 Each archegonium has a short, projecting neck with venter embeded. The jacket of neck is
one-cell thick and is made up of two tiers of four cells each. Cover cells/lid cells are absent.
 Antherozoids swim through rainwater or dew towards archegonium to fuse with egg forming a
diploid zygote that develops into embryo.
 The embryo is gradually pushed into the interior of female gametophyte (endoscopic embryo
development).
 In some species of Selaginella (e.g., S.rupestris) megaspore is not released from the
megasporangium. Even then fertilization and embryo development occurs while the
megaspore is still inside megasporangium (vivipary).

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Diagrammatic representation of Life Cycle of Selaginella

Fig 7. Selaginella Life-cycle.

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SOLVED PROBLEMS

Subjective

Prob 1. What do you understand by the terms “annulus” and “stomium”?

Sol. The annulus is a ring like portion of the jacket consisting of vertically elongated cells. It
helps in the dehiscence of sporangium and dispersal of spores. Stomiurn consists of group
of long, flat and thin walled cells. Stomium permit the early cleavage of the sporangium and
helps in it’s dehiscence.

Prob 2. What is ‘false induslum’ in Pteris? Mention its function.

Sol. In Pteris, each sorus is protected by reflexed margin of the fertile pennule, which, is called
false indusium. It is protective in function.

Prob 3. What are fronds? Give an example.

Sol. The foliage leaves of ferns are called fronds. e.g. : Pteris.

Prob 4. What are ramenta in Pteris?

Sol. Brown hair like structures present on the base of petiole and rhizome are cailea ramenta.

Prob 5. Name the vernation shown by young leaves of Pteris. Define.

Sol. The young leaves in Pteris are coiled like a watch spring and this condition is called
circinate vernation.

Prob 6. What is coenosorus In Pteris?

Sol. The sorus which is long, linear, continuous and formed by the fusion of several son is called
the coenosorus.

Prob 7. In which plant do you find mixed type of sorus? Define It.
Sol. Mixed sorus is present in Pteris.
A sorus consisting of older sporangium in the middle and younger sporangia to right and left
of older sporangium is called the mixed sorus.

Prob 8. Write briefly about dehiscence of sporangium in Pteris.

Sol. In Pteris, the dehiscence of mature sporangium takes place in dry atmosphere. The annulus
contracts and the sporangial wall ruptures transversely in the region of stomium.
The liberation of spores is by “Sting mechanism” or “Catapult mechanism”.

Prob 9. How many neck canal cells are present in the archegonium of Pteris ? In what way they
differ from those of Funana?
Sol. In the archegonium of Pteris, only one neck canal cell is present. It is binucleate. In Funaria,
neck canal cells are six or more and each cell is uninucleate.

Prob 10. Why self-fertilization is not carried out in the gametophyte or Pteris?
Sol. The gametophyte of Pteris is monoecious. The antheridia matures first (Protandry) cross
fertilization takes place. So self - fertilization is not carried out.

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Objective

Prob 1. “Botanical snakes” are

(A) Algae (B) Bryophytes
(C) Pteridophytes (D) Gymnosperms

Sol. (C). Pteridophytes are also called ‘Snakes of plant kingdom’ or ‘Botanical snakes’ as
snakes, i.e. reptiles evolved after amphibians.

Prob 2. Pteridophytes are characterized by

(A) Presence of vascular tissue (B) Absence of flowers
(C) Absence of fruits (D) All of the above

Sol. (D). Pteridophytes are vascular cryptogams.

Prob 3. Vessels are present in the xylem of

(A) Lycopodium (B) Selaginella
(C) Equisetum (D) Both B and C

Sol. (D). Vessels are usually absent in pteridophytes but exceptionally present in selaginella and
equisetum.

Prob 4. Megasporophyll of pteridophytes is comparable to which structure of angiosperms?

(A) Carpel (B) Microsporophyll
(C) Stamen (D) Ovule

Sol. (A). Megasporophyll in pteridophytes is associated with formation of megaspores and

hence is comparable to carpel of angiosperms.

Prob 5. Fern prothallus normally is

(A) Haploid (B) Diploid
(C) Triploid (D) Tetraploid

Sol. (A). Prothallus in fern is produced by germination of haploid spore and hence haploid.

Prob 6. Sprophyte of pteris is

(A) Independent of gametophyte from the very beginning
(B) Dependent upon gametophyte only in the beginning
(C) Dependent on gametophyte completely
(D) None of the above

Sol. (B). Sporophyte initially depends on gametophyte and later becomes independent in pteris.

Prob 7. In pteris, the sori are

(A) Continuous and linear (B) Discontinuous and circular
(C) Discontinuous and reniform (D) Discontinuous and vermiform

Sol. (A). Sporangia are grouped in continuous and linear type of sorus. The sorus is called as
Coenosorus.

Prob 8. Which of the following pteridophytes show the chloroplast having pyrenoids?
(A) Equisetum (B) Selaginella
(C) Pteris (D) Marsillea

Sol. (B). Pyrenoids in chloroplast which is an algae character are present in mesophyll cells of
sepaginella.

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Prob 9. A stele having leaf gaps to produce many smaller meristeles is known as
(A) Dictyostele (B) Solenostele
(C) Siphonostele (D) All of these

Sol. (A). In P.Vittata the stell is a ‘dictyostele’ with a ring of vascular strands which are called
meristeles.

Prob 10. In Selaginella the glossopodium is present on

(A) Root (B) Rhizophore
(C) Stem (D) Leaf

Sol. (D). The leaves are ligulate, at the base of ligule, a sheath of elongated cells called
glossopodium is present.

ASSIGNMENT

Subjective

Very short answer type questions

1. Mention the systematic position of Pteris.
2. What are fronds? Give an example.
3. Name the two species having unipinnate and imparipinnate leaves.
4. What are ramenta in Pteris.
5. Name the vernation shown by young leaves of Pteris? Define.
6. What is a dictyostele? Give an example.
7. Define ‘false indusium’ in Pteris? Mention its function.
8. What is coenosorus in Pteris?
9. In which plant do you find mixed type of sorus? Define it.
10. Distinguish between ‘annulus’ and ‘stomium’ of Pteris.
11. A prothallus of pteris has 10 archegonia and 10 antheridia. What is the total number of
sperms formed in the prothallus?

Short answer type questions

12. Describe briefly the external morphology of Pteris plant.
13. Write about sorus in Pteris.
14. Explain the structure and dehiscence of sporangium of Pteris.
15. Describe the structure of antheridia and archegonia of Pteris.
16. Differentiate between archegonia of Funaria and Pteris.
17. Describe the structure of Pteris prothallus.

Long answer type questions

18. Describe the internal structure of rhizome of Pteris.
19. Explain the method of reproduction in the sporophyte of Pteris.
20. Write about the process of reproduction in the gametophyte of Pteris.

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Objective

LEVEL – I
1. Members of pteridophytes are
(A) Homosporous (B) Heterosporous
(C) Homosporous or heterosporous (D) None of the above
2. Spores on germination in pteridophytes give rise to
(A) Sporophyte (B) Strobilus
(C) Prothallus (D) None of these
3. Which of the following is heterosporous?
(A) Selaginella (B) Salvinia
(C) Azolla (D) All of these
4. Sex organs in pteridophytes are
(A) Unicellular and jacketed (B) Multicellular and non-jacketed
(C) Multicellular and jacketed (D) Unicellular and non-jacketed
5. Number of neck canal cells in archegonium of pteridophytes is
(A) 2 - 6 (B) 4
(C) 1 (D) 1 - 14
6. The spermatozoids in vascular cryptogams are
(A) Non-flagellated (B) Biflagellated
(C) Multiflagellated (D) Biflagellated or multiflagellated
7. The characteristic feature of eusporangiate type of sporangium development is
(A) Sporangium develops from group of cells (B) Having multi-layered jacket
(C) Contain large number of spores (D) All of the above
8. ‘Stele’ includes
(A) Vascular tissue (xylem and phloem)
(B) Pith and vascular tissue
(C) Vascular tissue + pith + pericycle
(D) Vascular tissue + pith + pericycle + endodermis
9. Most primitive type of stele found in pteridophytes is
(A) Siphonostele (B) Protostele
(C) Solenostele (D) Dictyostele
10. Protostele with pith or medullated protostele is called
(A) Solenostele (B) Siphonostele
(C) Actinostele (D) Plectostele
11. Indusium in Pteridium is:
(A) Outer true and inner false (B) Only true
(C) Outer false and inner true (D) Only false
12. Spores of fern are:
(A) Haploid (B) Diploid
(C) Triploid (D) Polyploid
13. Sporophyte of Pteridium is:
(A) Dependent on gametophyte (B) Dependent on embryo
(C) Dependent on prothallus (D) Independent
14. Ligule in leaf of Selaginella is present on surface:
(A) Abaxial (B) Adaxial
(C) Abaxial or adaxial (D) None of these
15. What is produced from the fertilization of egg in fern?
(A) Ascospore (B) Zygospore
(C) Oospore (D) None of these
16. Which is epiphytic sps. of Selaginella?
(A) Selaginella kraussiana (B) S. chrysocaulous
(C) S. oregena (D) All of these
17. In which pteridophyte, apospory was first observed?
(A) Lycopodium (B) Isoetes
(C) Selaginella (D) Athyrium
18. Fern plant is a:
(A) Haploid gametophyte (B) Diploid gametophyte
(C) Diploid sporophyte (D) Haploid sporophyte

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19. The stem of Selaginella is automatically characterised by the presence of:

(A) Siphonostele (B) Amphiphloic siphonostele
(C) Ectophloic siphonostele (D) Protostele
20. A plant having vascular supply, producing spores but lacking seeds is a:
(A) Bryophyte (B) Pteridophyte
(C) Gymnosperm (D) Angiosperm

LEVEL – II
1. Afernisa:
(A) Vascular non-flowering plant (B) Non-vascular non-flowering plant
(C) Vascular flowering plant (D) Non-vascular flowering plant
2. If the number of chromosomes in the foot of a fern embryo is 8, what should be the number in
its spore?
(A) 4 (B) 8
(C) 16 (D) 23
3. The cells of fern prothallus contain nucleus with:
(A) 4-n chromosomes (B) 3-n chromosomes
(C) 2-n chromosomes (D) n chromosomes
4. The first seed plants appeared during which period?
(A) Silurian (B) Devonian
(C) Carboniferous (D) Cretaceous
5. Seed habit first originated in:
(A) Certain ferns (B) Certain pines
(C) Certain monocots (D) Certain dicots
6. Which of the following plants exhibit independent alternation of generation?
(A) Angiosperms (B) Gymnosperms
(C) Pteridophytes (D) Bryophytes
7. In which of the following rocks the earlier vascular plants have been discovered?
(A) Early cambrian (B) Early devonian
(C) Mid cretaceous (D) Silurian
8. While entering the neck of a fern archegonium, sperms show:
(A) Phototaxy (B) Chemotaxy
(C) Thermotaxy (D) Cyclosis
9. Chlorophyll and chloroplast are present, in spores of:
(A) Lycopodium (B) Chemotaxy
(C) Dryopteris (D) Marsillea
10. Which of the following pteridophytes shows the chloroplast having pyrenoids?
(A) Equisetum (B) Selaginella
(C) Pteridium (D) Marsillea
11. Which of the following has amphiphloic siphonostele?
(A) Rhizome of Marsillea (B) Stem of Lycopodium
(C) Rhizome ofPteris (D) Stem of Equisetum
12. Heterospory and seed habit are often discussed in relation to a structure called:
(A) Spathe (B) Bract
(C) Petiole (D) Ligule
13. Botanical name of Sanjeevani or Hasnpari is:
(A) Selaginella bryopteris (B) S. chrysocaulous
(C) S. kraussiana (D) S. pallidissima
14. ‘Maiden hair fern’ is:
(A) Dryopteris (B) Azolla
(C) A diantum (D) Pteris
15. The rudimentary seed habit has been attained in:
(A) Psilotum (B) Lycopodium
(C) Selaginella (D) Equisetum
16. Main plant of Selaginella species is:
(A) Gametophyte (B) Sporophyte
(C) Both (A) and (B) (D) Halophytes
17. Sporocarp is a reproductive structure of:
(A) Some algae (B) Some aquatic ferns having son
(C) Angiosperms having spores (D) Bryophytes

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18. Protostele is:

(A) Stele divided into many parts
(B) Early stele with central solid xylem surrounded by phloem
(C) Latent stele
(D) Angiosperm stele
19. Polystelic condition is found in:
(A) Rachis of Pteris (B) Roots of Dryopteris
(C) Stem of all species of Selaginella (D) Stem of Selaginella willdenowii
20. The vascular supply given from the main stele for leaf is called:
(A) Leaf gap (B) Leaftrace
(C) Branch trace (D) 1

LEVEL – III
1. If a sporangium is derived from a single cell, it is called:
(A) Leptosporangiate (B) Eusporangiate
(C) Heterosporangiate (D) None of these
2. Indusium is found in:
(A) Fungi (B) Moss
(C) Algae (D) Pieris
3. The walking fern is so named because:
(A) Its spores are able to walk
(B) It is dispersed through the agency of walking animals
(C) It propagates vegetatively by its leaf tips
(D) It knows how to walk by itself
4. Prothallus means:
(A) immature gametophyte (B) Immature sporophyte
(C) Immature archegonium (D) None of these
5. Which one of the following is the earliest land plant?
(A) Cordaites (B) Cooksonia
(C) Hornea (D) Rhynia
6. In ferns, meiosis occurs during:
(A) Spore formation (B) Gamete formation
(C) Antheridia formation (D) All of these
7. A pteridophyte which fixes N2 is:
(A) Azolla (B) Salvinia
(C) Pteris (D) Selaginella
8. In ferns, archegonia are found on:
(A) Prothallus (B) Leaves
(C) Stem (D) Sporangia
9. Large leaves of ferns are called fronds which are:
(A) Reproductive (B) Vegetative
(C) Foliage (D) All of these
10. Pteridophytes are characterized by:
(A) Dominant sporophytic generation
(B) Dominant gametophytic generation
(C) Formation of leafs’ gametophore as dominant generation
(D) None of the above
11. Pteridophytes differ from bryophytes in having:
(A) Independent sporophyte (B) Dependent sporophyte
(C) No vascular bundles (D) None of the above
12. Structure present over the leaves of ferns is called:
(A) Ramenta (B) Fronds
(C) Spathe (D) Indusium
13. Coal is formed by:
(A) Pteridophytes (B) Bryophytes
(C) Fungi (D) Bacteria

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14. The 13-celled male gametophyte in Selaginella is:

(A) 12-cells of antheridium + 1-prothallial cell
(B) 10-cells of antheridiurn + 3-prothallial cells
(C) 9-cells of antheridium + 4-prothallial cells
(D) 7-cells of antheridium + 6-prothallial cells
15. Sometimes the prothallus of a fern gives rise to a fern plant without any fertilization. This is an
example of:
(A) Apospory (B) Apogamy
(C) Parthenocarpy (D) Parthenogenesis
16. Polystelic condition is found in:
(A) Rachis of Pteris (B) Roots of Dry pteris
(C) Stem of all species of Riccia (D) Stem of Selaginella laevigala
17. Vascular strands in rhizome of Pteridium are:
(A) Collateral open (B) Bicollateral open
(C) Concentric amphivasal (D) Concentric amphicribal
18. ‘Club moss’ is the common name of:
(A) Lycopodium (B) Selaginella
(C) Funaria (D) Potanogeron
19. In fern, gametophyte is:
(A) Main plant body
(B) Prothallus attached to the sporophyte
(C) Prothallus free from sporophyte
(D) A structure attached to the rhizome
20. In Selaginella, a member of Pteridophytes, heterosporous spores are:
(A) Large and small (B) Hap bid and diploid
(C) Asexual and sexual (D) All spores of same size

ANSWERS TO ASSIGNMENT
Objective
Level  I
1) C 2) C 3) D 4) C 5) D
6) D 7) D 8) C 9) B 10) B
11) C 12) A 13) D 14) B 15) C
16) D 17) D 18) C 19) D 20) B

Level  II
1) A 2) A 3) D 4) C 5) A
6) C 7) D 8) B 9) A 10) B
11) A 12) D 13) A 14) C 15) C
16) B 17) B 18) B 19) D 20) B

Level  III
1) A 2) D 3) C 4) A 5) B
6) A 7) A 8) A 9) D 10) A
11) A 12) A 13) A 14) A 15) B
16) D 17) D 18) A 19) C 20) A

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