 Some floral parts like calyx and various
Sexual Reproduction in
Flowering Plants
STRUCTURE OF FLOWER
NON-ESSENTIAL / ACCESSORY WHORLS
 Calyx: Formed of green sepals ; (Outermost)
 Corolla: Formed of colored or white petals ; (Inner
to calyx)
ESSENTIAL / REPRODUCTIVE WHORLS
 Androecium: The male reproductive whorl formed
of stamens (microsporophylls) ; (Between Corolla
and Gynoecium).
 Gynoecium: The female reproductive whorl
formed of carpels (megasporophylls) ;( Innermost).
Sexual reproduction is the process of development of new
organisms from two parents through fusion of male and
female gametes.
The sporophylls (reproductive organs) are of two types-
 Megasporophylls (Carpel) is distinguished as
ovary bearing ovule, style and stigma.
 Microsporophylls (Stamen) is distinguished as
filament, anther and connective.
In flowering plants, the reproductive phase is marked by the
initiation of flower development.
During the reproductive season, several hormonal and
structural changes in the apices of twigs initiate
development of floral primordia.
Soon, inflorescences with floral primordia are formed.
Floral primordia develop first into floral buds and then into
flowers.
Floriculture is the branch of ornamental horticulture related
with growing and marketing of flowers and ornamental
plants.
FUNCTIONS OF A FLOWER
 Modified shoot meant to perform the function of
sexual reproduction.
 Shaped variously to assist in pollination, the
prerequisite for reproduction.
 The ovary gets transformed into fruit and ovules
into seeds after fertilization.
modifications in ovaries help the dispersal of fruits
and seeds.
FLOWER IS A MODIFIED SHOOT
According to Goethe, flower is a modified shoot, which is
adapted for reproductive process. This is evident from the
fact, that the thalamus (base) of many flowers bear small
nodes and internodes.
In some plants when an internode between corolla and
androecium become longer in size, it is called androphore.
Ex- Passiflora
In some plants when an internode between androecium and
Gynoecium becomes longer in size. This is called
gynophore. Ex- Capparis
In some other plants when an internode between calyx and
corolla becomes longer in size. This is called anthophore.
Ex-Dianthus
TYPES OF FLOWERS
Essential and non-essential whorls- Gynoecium and
androecium are called essential whorls because these are
directly involved in the process of sexual reproduction. A
flower must contain at least one of the two whorls. The
calyx and the corolla are referred to as the non-essential
whorls, because they are not responsible for the formation of
gametes and seeds.
Unisexual (Imperfect)- When only one whorl is present.
(When only androecium is present, flower is called
staminate, and when only Gynoecium is present, the flower
is called pistillate.)
Bisexual or Hermaphrodite (Perfect)- When both
essential whorls are present in the same flower.
Monoecious- An individual plant, bearing unisexual flower
of both sexes separately i.e., staminate and pistillate flowers
are present on same plant.
Dioecious- A plant containing flowers of only one sex. The
term, dioecious, is used when a species has separate
staminate-flowered plants and pistillate-flowered plants.
Complete flower- When all the four whorls are present in
the flower, it is called complete flower.
Incomplete flower- If any of the non-essential whorls is not
present, the flower is incomplete.
Neuter flower- When both of the essential whorls are
absent.
Isomerous flower- When each whorl of a flower has equal
number of units.
Heteromerous flower- When each whorl of a flower has
unequal number of units.
Hypogynous flower- When the ovary of the pistil is above
the site of attachment of other floral parts (stamens, sepals,
petals), it is called a superior ovary. Flower with such
ovary are called hypogynous flower.
Perigynous flower- These are flowers show varying
degrees of ovary positions from partially superior to
partially inferior.
Epigynous flower- When the ovary of the pistil is below the
site of attachment of other floral parts (stamens, sepals,
petals), it is called a inferior ovary. Flower with such ovary
are called epigynous flower.
Lower plants like Bryophytes-Gametophytic phase is
dominant.
Higher plants like pteridophytes, gymnosperms and
angiosperms- sporophytic phase is dominant and chief.
. In angiosperms Gametophytic phase is highly reduced and
microscopic.

SPOROPHYTIC PHASE
 It is the diploid phase (always).
 It is developed from diploid zygote. Therefore
zygote represents the first cell of sporophytic phase
as it gives rise to spores.
 Spores are haploid cells but not gametes.
 A sporophyte (since it is diploid) produces haploid
spores with the help of meiosis (also called
reductional division).
Anthesis is the term used for the time when a flower comes
into full bloom. It is a time when the flower is fully In higher plants, like angiosperms, spores are of two types.
expanded.
Male plants produce small sized spores, called microspores
VEGETATIVE SEXUAL REPRODUCTION (Or pollen grains).
REPRODUCTION
No specialized structures are Specialized structures called Whereas the female plant (part) produces larger spores,
used. gametes are used. called megaspores (macrospores).
No fertilization. Fertilization essentially occurs.
 In angiosperms, the entire standing plant body is
The ploidy of the cells The ploidy of the cells changes sporophytic (diploid).
remains same throughout the alternatively from diploidy to  Gametophytic tissues are highly reduced and
process. haploidy and again to diploidy. develop inside the small part of flower.
 Microspores are developed inside microsporangia
(anthers), while megaspores develop inside
Megasporangium (ovule), within ovaries.
Offspring are similar to Offspring differ in some of
parents (natural clones). their characters from their GAMETOPHYTIC PHASE
parent plants. (Hybrids).
 It is the haploid phase of the plant.
 It is a very reduced phase in higher plants like
The whole process of sexual reproduction in flowering angiosperms, but is very important.
plants can be divided into three steps:-  It develops from spores (which are haploid
structures). Therefore, spores represent the first
cells of gametophytic phase.

Since there are two types of spores — microspores (pollen

1. Pre-fertilization Events grains or male spores) and megaspores (or female spores),
 Gametogenesis the gametophytes are also of two types — male
→ Microsporangium and gametophyte (it develops from microspore) and female
microsporogenesis (Pollen Grain gametophyte (it develops from megaspore).
formation) (male gametes)
→Megasporangium and  Gametophytes on development give rise to
megasporogenesis (Embryo sac gametes.
formation) (female gametes  Male gametophyte produces male gametes
 Pollination (sperms), while female gametophyte produces
2. Fertilisation female gametes (egg cell or ovum).
3. Post-fertilization Events  In angiosperms, the mature male gametophyte
 Endosperm formation consists only of 3 cells (It contains one tube cells
 Embryogenesis and 2 sperms).
 Development of seed and fruit  Its development starts when the microspores are
within the anther but gets completed only when it is
In all categories of higher plants, 2 distinct phases are found transferred to the stigma of the flower (through
in the life cycle which regularly alternate with each other. pollination).
 The mature female gametophyte in angiosperms is
 Sporophytic phase mostly 7 celled (but having 8 nuclei).
 Gametophytic phase
  It is also called as embryo sac. It contains, an egg
cell, 2 synergid cells, 3 antipodal cells and one
large embryo sac cell.
 The gametes from two types of gametophytes unite
(fertilise) to produce a diploid cell, called zygote,
which sets seeds and again starts the sporophytic
phase.
 In this way, a regular alternation between
sporophytic and gametophytic stage is clearly
apparent in angiosperms.
ACCESSORY WHORLS
CALYX
1. Outermost whorl of a flower.
2. Consists of sepals which are usually green.
3. Sepals may be free from each other (polysepalous)
or fused (gamosepalous).
4. Encloses and protects the inner whorls in the bud
stage.
5. Since sepals contain chlorophyll, they can also
synthesize food.
COROLLA
1. Found on the inside of the calyx and is the most
conspicuous part.
2. Usually white or brightly colored.
3. The brightly colored corolla attracts agents of
pollination such as birds and insects.
4. Units are called petals (much larger than sepals).
The petals may be separate from each other
(polypetalous) or become partly or completely
fused (gamopetalous).
5. If the petals are completely fused, they form corolla
tube.
6. Calyx and corolla together-Perianth
7. Units of Perianth- Tepals
MALE REPRODUCTIVE
STRUCTURE
1. STAMENS / ANDROECIUM
1. Found inner to the corolla and are attached either to
the thalamus or to the petals.
2. The stamens or microsporophylls collectively form
the male whorl or androecium.
3. The number, length and arrangement of stamens
vary in flowers of different species. Each stamen
has following three parts-
A. Filament: It is the stalk-like part of a stamen that
supports anther terminally in the most suitable position for
pollen transfer to take place, and is attached to the thalamus
by its basal end.
B. Anther: (Microsporangia) produce microspores (pollen
grains) which produce male gametophyte. It is the swollen
bilobed structure at the terminal end of a filament which
consists of two anther lobes (theca). Hence are called
dithecous.
An anther with two anther lobes is said to be dithecous.
Majority Of plants have dithecous anthers.
Exceptions- Malvaceae (Hibiscus) & Euphorbiaceae
(Euphorbia). Here each anther has only one lobe
(monothecous).
Each anther lobe has two pollen sacs or pollen chambers
which are demarcated by a longitudinal groove on the
ventral surface. This denotes the line of dehiscence of anther
lobes to liberate pollen grains formed in the pollen sacs.
A dithecous anther, therefore, has four pollen sacs and is
called tetrasporangiate, while monothecous anther is
bisporangiate.
C. Connective: Extension of a filament between two anther
lobes to join them together.
In some plants, one
or more stamens are
nonfunctional and
without anther, as in
roses. Such
nonfunctional
stamens are called
staminodes.
2. MICROSPORANGIA (SING.
MICROSPORANGIUM) / POLLEN SACS
The pollen chambers represent microsporangia. Thus, each
anther consists of four microsporangia, two in each anther
lobe. These are covered by the common epidermis of anther
which contain pollen grains or microspores.
DEVELOPMENT OF MICROSPORANGIUM
A very young anther consists of a group of actively dividing
parenchymatous cells surrounded by layer of epidermis.
It soon becomes two-lobed. Each anther lobe develops into
two microsporangia as rounded mass of archegonial cells.
The four microsporangia of an anther finally develop into
four pollen sacs. Development of microsporangia is
eusporangiate type (from a group of initial cells).
Archegonial cells of each microsporangium divide by
periclinal divisions to give rise to an outer primary parietal
layer and an inner primary sporogenous layer. The primary
parietal layer divides to form concentric layers of pollen sac
wall and the primary sporogenous layer divides to produce a
mass of sporogenous cells or microspore mother cells.
They are initially connected by plasmodesmata. They are
broken by the formation of callose layers inner to cell wall
and microspore mother cells separate from each other.
STRUCTURE OF MICROSPORANGIUM
1. Few cells in the hypodermal region become differentiated
as archesporial cells. In Boerhaavia and Dionaea, there is
only one archesporial cell.
2. The archesporial cell divides periclinally (along the
periphery) to form outer-primary parietal layer and inner-
sporogenous layer.

3. The primary parietal layer lies just beneath the epidermis

and divides again periclinally to form 3-5 concentric layers.
These layers give rise to the wall of the sporangium, along
with epidermis. In a transverse section, a typical
microsporangium appears circular in outline. Its wall has
four layers-

1. Epidermis: It is the outermost and one-cell thick layer.

2. Endothecium: It lies inside the epidermis. It is a single

layer of radially elongated cells with fibrous thickenings
which help dehiscence of anther. The cells of endothecium
in the shallow groove between two microsporangia remain
thin-walled and form stomium.
3. Middle Layers: These are three to four layers of thin-
walled cells which disintegrate in mature anthers.These
three layers protect sporogenous cells and help in
dehiscence of anther to release pollen.
4. Tapetum: This is the innermost layer of cells with dense
cytoplasm. Tapetum nourishes the developing pollen grains
(microspores) and secretes both enzymes and hormones. Its
cells have more than one nuclei. These cells are of following
two types:
Amoeboid or Invasive or Periplasmodial cells grow and
form a periplasmodium that provides nourishment and other
materials to spore mother cells, e.g., Lily, Tradescantia,
Typha, Alisma, etc.
Secretory or Glandular or Parietal cells provide
nourishment to sporogenous cells, e.g., Symphoricapus.
In the end, both the cell types of tapetum degenerate.
FUNCTIONS OF TAPETUM
1. Provides nourishment to the growing sporogenous
cells, microspore mother cells and growing
microspores.
2. Secretes enzymes and hormones (IAA) that are
stored in pollen grains for their early growth.
3. Secretes enzyme callase for the dissolution of
callose binding the microspores.
4. Produces Ubisch granules that provide
sporopollenin and other materials for exine part of
pollen grain covering.
5. Provides sporopollenin covering around
entomophilous pollen grains which provide
resistance against physical and biological
destructive forces, therefore, pollen can be
preserved for a long time.
6. Pollenkitt and tryphine are transferred by tapetum
to pollen surface.
7. Secretes special proteins for pollen grains to
recognize compatibility-incompatibility.
Formation of Microspores or Pollen Grains
Formation of microspores or pollen grains from the
microspore mother cells or pollen mother cells (PMC) or
primary sporogenous cells occurs inside microsporangia and
is called microsporogenesis.
(i) The sporogenous layer, formed by archesporial cells
divides many times to form pollen mother cells (or
microspore mother cells). These are diploid cells.
(ii) The microspore mother cells remain surrounded by a
wall of callose while they are dividing by meiosis. Because
of its highly impermeable nature, callose provides an
isolation barrier separating the microspore mother cells from
each other. It prevents the microspores from sticking with
each other.
(iii) Each such cell divides by meiosis and forms four
haploid microspores or pollen grains.
(iv) The division is of two types in various angiosperms -
simultaneous type and successive type.
(v) In simultaneous division, the M1 of meiosis is not
followed by cytokinesis. Wall formation takes place only
after the completion of both M1 and M2.
(vi) While in successive type of division, cytokinesis occurs
after both divisions M1 as well as M2.
Each microspore mother cell undergoes meiosis and gives
rise to four haploid microspores. The microspores are
arranged in a cluster of four cells and form the microspore
tetrad. There are five types of tetrads-
As the anthers mature and dehydrate, microspores dissociate
from each other and form pollen grains.
COMPOUND POLLEN GRAINS AND POLLINIUM:
In Typha, Drosera, Juncus, etc., all the four pollen grains of
tetrad do not separate but together form compound pollen
grains. In Mimosoideae, 8-64 pollen grains may remain
united.
In Calotropis and many orchids, all the pollen grains of a
sporangium form a single mass called pollinium. The two
adjacent pollinia are connected to a sticky corpusculum to
form a translator.

DEHISCENCE OF ANTHER
As the anther matures, it dries up. The strip between the two
pollen sacs of each anther lobe disintegrates and a single
cavity is formed. With the loss of water, the dead cells of
endothecium contract from their outer thin walls. As the
outer radial walls come nearer, endothecium shortens and
the anther lobe wall ruptures in the region of stomium. The
exposed spores are dispersed by various agencies of
pollination.
STRUCTURE OF MICROSPORES OR POLLEN
GRAINS
 They are male gametophytes.
 They are usually unicellular, spherical,
and haploid measuring about 25-50
micrometers in diameter.
 They may be oval, ellipsoidal, triangular,
lobed or even crescent-shaped.
 The outer surface of pollen grains may
have spines, ridges or furrows which may
vary in different species.
The germ pores are the points from where the pollen tube
emerges out.
Monocolpate pollen grains have single furrow or pore for
the emergence of pollen tube. They are found in family
liliaceae and monocots.
Tricolpate pollen grains have three furrows or three rows of
pores through which pollen content comes out. They are
found in true dicots.
Each pollen grain (Microspore) represents the first cell of
male gametophyte. The wall or covering of the pollen grain
is called sporoderm. It consists two layers, the outer exine
and inner intine.
EXINE
It is thick and ornamented. It is chiefly composed of
sporopollenin, the most resistant organic material known. It
can withstand high temperature, strong acids and alkali and
cannot be degraded by any known enzyme. Pollen grains are
well preserved in nature as fossils because of the presence of
sporopollenin. Exine also possesses proteins for enzymatic
and compatibility reactions.
The exine consists of two layers - outer ektexine (sexine)
and inner endexine (nexine). The ektexine further
comprises an inner continuous foot layer, a middle
discontinuous baculate layer and an outermost
discontinuous tectum. Tectum has characteristic
sculpturing. It helps in identifying the pollen grains and
referring them to their family, genus or species.
The study of external morphology of mature pollen grains is
called palynology.
INTINE
It is thin and uniform. It is made up of pecto-cellulose
(pectin+cellulose). At the time of pollen germination, intine
comes out of the germ pore in the form of a pollen tube. At
places, intine contains enzymatic proteins.
POLLEN ALLERGY
In some people, the exposure to pollen grains cause severe
allergies and bronchial afflictions often leading to chronic
respiratory disorders, such as asthma, bronchitis, etc. This
type of allergic reaction occurs mainly due to anemophilous
pollens, which fly with wind current. One best example of
pollen allergy occurs with the pollen of Parthenium or
carrot grass. It is not a native plant, but was imported
earlier along with wheat varieties.
Now, it has spread to several places and causes pollen
allergy and hay fever, which are caused by a poisonous
substance, tryphine. This is composed of poisonous
proteins and hydrophilic substances.
POLLEN PRODUCTS
Pollen grains are rich in nutrients. They are given as tablets
or syrups to athletes and race horses to enhance their
performance.
POLLEN VIABILITY
It is the period, during which pollen retains its ability to
germinate. After dehiscence (shedding), pollen must be
landed over the compatible stigma, before they lose their
viability. Viability of pollens varies greatly in different
plants. For example it is about 30 minutes in wheat and rice,
but months and years in many other (members of families
Rosaceae, Leguminoseae and Solanaceae). Environmental
conditions and temperature play an important role in
maintaining viability of the pollen grains.
POLLEN BANKS
These days pollen grains of a large number of species can be
stored for years in liquid nitrogen (−196∘C). Such stored
pollen can be used as pollen banks like the seed banks stored
for crop breeding experiments.
DEVELOPMENT OF MALE GAMETOPHYTE
Development of male gametophyte starts when the pollens
are within the anther lobes. Thus initial development of
male gametophyte takes place inside the anther lobes
(microsporangia). But when half of the germination (i.e.,
upto 2-3 cell stage) gets completed, the anther dehisces and
the germinating pollen (male gametophytes) are shed. Now
pollination takes place (ie., these pollens are transferred to
the stigma of the carpel of a flower). Further development of
male gametophyte takes place on the stigma.
In this way the development of the male gametophyte
involves following steps:
 Development of male gametophyte before
pollination.
 Dehiscence of anther and liberation of
developing pollens
 Pollination
 Development after pollination.
(A) DEVELOPMENT OF MALE GAMETOPHYTE
BEFORE POLLINATION
(i) Development of male gametophyte before pollination
occurs inside the anthers (microsporangia).
(ii) During germination, the nucleus of the microspore is
displaced from centre to one side. This displacement is
always in a particular direction and marks the position of the
germinative cell.
(iii) The cytoplasm becomes highly vacuolated.
(iv) Now microspore undergoes mitotic division and forms
two unequal cells (1) the larger cell, called vegetative cell
and (ii) the smaller cell, called generative cell.
(v) The vegetative cell is bigger, has a large amount food
reserve and a large irregularly shaped nucleus.
(vi) The generative cell is small. It floats in the cytoplasm of
the vegetative cell. It is spindle shaped with dense
cytoplasm and a nucleus.
(vii) The vegetative cell on germination gives rise to pollen
tube (after pollination, on stigma).
(viii) The generative cell produces two male gametes by one
more mitosis (after pollination).
(ix) In about 60% angiosperms, the pollens are shed (fall) at
two-cell stage (vegetative and generative cell stage), and
further development of the male gametophyte takes place on
the stigma, after pollination. In some cases, the pollen are
shed at three celled stage (vegetative cell and two male
gametes). Thus, in these plants, the gamete formation takes
place before pollination.
(B) DEHISCENCE OF ANTHER AND LIBERATION
OF DEVELOPING POLLENS
When pollens get matured (usually at 2 celled stage), they
exert some pressure on anther wall. Consequently, the wall
of the anther bursts and the germinating pollen grains are
released. This is called dehiscence of anther.
(C) POLLINATION
Transfer of pollen grains from anther to the stigma (of a
pistil) is called pollination. It is of two main types- (i)
Self-pollination (autogamy and geitonogamy) (ii)
Cross-pollination (Xenogamy).
1. SELF POLLINATION
Transfer of pollen from anther of a flower to the stigma of
the same flower (or flower of same plant) is called self
pollination.
Self-pollination can be of two types, autogamy and
geitonogamy.
(A) AUTOGAMY: It is a type of self-pollination that is
found only in bisexual flower. In this case, the stigma of a
flower is pollinated by its own pollen. Autogamy occurs by
three methods.
(i) Homogamy: The anthers and stigmas of open
flowers mature at the same time, e.g., Catharanthus (Vinca),
Mirabilis (Four O'Clock plant).
(ii) Cleistogamy: The flowers remain closed
during pollination (with the help of its petals). Thus, stigma
receives only the pollen of its own flower, e.g., Oxalis,
Viola etc.
(iii) Bud Pollination: In this case the pollination
occurs in the bud stage. The sex organs develop before the
opening of bud, thus internal pollination takes place, e.g.,
Pea, Wheat, Rice etc.
(B) GEITONOGAMY : In this type of pollination, the
pollen grains of one flower are transferred to the stigma of
another flower belonging to either the same plant or
genetically similar plant. This type of pollination looks
similar Lo a cross pollination as it involves a pollinating
agency. But in fact, it is treated as self pollination because,
pollen and stigma belong to the same plant and therefore
there is no genetic differentiation.
So, geitonogamy is functionally a cross-pollination
involving a pollinating agent, but genetically it is similar to
autogamy since the pollen grains come from the same plant.
CHASMOGAMOUS AND CLEISTOGAMOUS
FLOWERS
Some plants like Viola (common pansy), Oxalis and
Commelina produce two types of flowers - (a)
Chasmogamous and (b) Cleistogamous.
In Chasmogamous flowers, the anther and stigma are fully Most of the plants are chasmogamous type (i.e., they
exposed to the external pollinating agencies, so, always expose their anthers and stigma to the pollinating agencies).
cross pollination occurs. As the pollen grains do not have locomotory structures, they
But cleistogamous flowers, remain closed and do not open are transferred from anther to the stigma, with the help of
at all. So, always self pollination occurs. certain agencies, called as pollinating agencies. Some of
ADVANTAGES OF SELF POLLINATION them are wind (air), water, insects, bats, birds and even man.
(1) It maintains purity of the species, by preserving all the Pollinating agencies may be biotic or abiotic. Two important
parental characters. abiotic agencies are wind and water. Pollen grains coming in
(ii) It is used to obtain pure-line characters (homozygosity) contact with wind/water is a chance factor. So, such plants
during breeding experiments. which depend upon wind/water, produce enormous amount
(ii) Plant does not have to depend on pollinating agencies. of pollens.
(iv) Only a small number of pollen grains are required. Further, majority of the plants use biotic agencies. Of all the
(v) Self-pollination strengthens the better characters of the biotic agencies insects are the most important.
plant. 1. ANEMOPHILY
DISADVANTAGES OF SELF POLLINATION Anemophily refers to the pollination by wind. Wind
(i) It does not eliminate bad characters from the race. pollinated flowers (a primitive feature) show following
(ii) Vigour and vitality of the race decreases, as there is no characters :
hybrid vigour.  Flowers are inconspicuous and not showy. They
(iii) Immunity to diseases decreases. are devoid of fragrance and nectar.
(iv) Ability to adapt according to changing environment  They produce a very large amount of pollen grains,
decreases. as considerable amount of pollen never reaches the
proper stigma.
2. CROSS POLLINATION (XENOGAMY OR  Anthers are versatile, hanging freely in air.
ALLOGAMY)  Pollen grains are dry, light and smooth walled.
Transfer of pollen from the anther of a flower to the stigma For effective wind pollination pollen grains should be
of another flower, belonging to a different plant is called light and non-sticky so that they can easily be
cross-pollination. It is usually found in unisexual flowers. transported by wind currents. These pollen grains have
However, it may also occur in bisexual plants as well. This well-exposed stamens (so that the pollens are easily
type of pollination helps in the formation of genetically dispersed into wind currents).
different plants, e.g., Hemp, Willow etc.  The flowers in wind pollinated plants have large
and feathery stigma to easily trap air-borne pollen
ADVANTAGES OF CROSS POLLINATION grains.
(i) It helps to eliminate bad characters from the race.  Wind pollinated flowers often have a single ovule
(ii) It helps in the development of new characters due to in each of their ovaries. Many flowers remain
recombination of genes. packed together in an inflorescence. A good
(iii) Vigour and vitality of the race increases, due to example of such inflorescence is found in corn cob.
heterosis. It is found commonly in grasses, most of the
(iv) Immunity to diseases increases. cereals and palms etc.
(v) Ability to adapt according to changing environment 2. HYDROPHILY
increases. Pollination through water current. Water pollinated flowers
AUTOGAMY GEITONOGAM XENOGAMY/
show following characters- Aquatic nature of the plant is no
Y ALLOGAMY
identification of its being pollinated through water. Many
It refers to self It also refers to It refers to cross
hydrophytes are pollinated through wind or insects (water
pollination. self pollination. pollination,
hyacinth and water lily). When it is with the help of water, it
Flower is always Flower may be
may take place completely under water (hypohydrophily) or
bisexual. unisexual or
may takes place on the water surface (epihydrophily).
Only one flower bisexual.
Pollination by water is not very common. It is found in
is used.
about 30 genera only, mostly monocots.
Transfer of Transfer of Transfer of pollen
Hydrophilous plants, like anemophilous flowers are
pollen from pollen from from anther to the
characterised by floral envelopes which are highly reduced
anther to the anther to the stigma of different
or even absent. It is commonly found in plants like Hydrilla,
stigma of same stigma of flower of different
Zostera (marine grass), Ceratophyllum and Vallisneria. In
flower. different flowers plant.
Vallisneria, the female flower reaches the surface of the
of same
water by its long stalk. The pollen grains from the male
plant.
flower are released into the water current and they finally
Flower is always Flower may be Flower may be reach the female flower. Water pollinated flowers show
bisexual. unisexual or unisexual of following features: → Water pollinated flowers are not very
bisexual. bisexual. colourful or bright.
Only one flower Two different Two different Also they have no fragrance (nectar).
is used. flowers of same flowers are used, Their pollen grains are buoyant, so that they can easily float.
plant which are ISC Biology-XII
are used which genetically Fig. 2.13: Water pollinated plant
are genetically different. • The pollen grains are coated with mucilage to protect
similar. themselves from getting wet. The stigma is large and
VARIOUS AGENCIES OF POLLINATION feathery to trap the pollen grains.
3. Entomophily
• Insects are the chief pollinators and it has been noted that
the evolution of flowering plant has gone hand-in-hand with
the evolution of the insects. Insects helping in pollination
are bees, flies, beetles, wasp, butterflies, ants and moths. Of
these, the first three are diurnal visiting flowers which opens
in day time and moths are nocturnal i.e. they visit flowers
which open after sunset.
Insect pollinated flowers show following characters:
* Flowers are brightly coloured.
◆ Pollen grains are sticky with a rough surface, so that
they may easily stick to Fig. 2.14: Pollination by insect
insect limbs.
→ Stigma is also sticky.
• A sugary fluid called nectar or honey is secreted in many
plants. In fact, insects visit flowers for nectar. Nectar glands
may be situated on thalamus, sepals, petals, carpels.
→ In some entomophilous flowers, special mechanisms are
seen. Night blooming flowers have white colour and
fragrance (so that they may be visible to insects).
In Centaurea (compositae), pistil bends and exposes the
stigma on being touched by insects. In Salvia, versatile
anthers and other balancing features, help in dusting of
insects with pollen.
• In case of Amorphophallus (tallest flower, about 6 feet),
insect lay their egg in the flowers and eventually carry out
pollination. In Yucca, also, the pollination is carried out by
a moth. Moth lays its egg in the locules of the ovary and
eventually the flower gets pollinated.
4. Ornithophily
Bird pollination is not so common. Humming bird, Sun
birds and honey eaters are some of the birds which visit
flowers and bring about pollination. Such flowers are
tubular, cup- shaped or urn shaped, bright in colour and
produce large quantities of pollen and plenty of nectar.
5. Chiropterophily
In this case the pollination is carried out by bats.
Chiropterophilous plants have flowers borne singly or in
clusters quite away from branches and leaves due to their
long stalk. These flowers open only at or after dusk. On
blooming, flowers emit an odour and produce large
quantities of nectar. Flowers are dull in colour. Bats, being
nocturnal are attracted by the odour of the flowers.
6. Malacophily
Fig. 2.15: Pollination by bird
Snails and slugs visit certain flowers and may be playing a
role in their pollination. This is generally observed in plants
like Arisaema (cobra plant) and also in arum lilies.
Outbreeding Devices (Efforts to Avoid Self Fertilisation)
35
Majority of flowering plants produce bisexual
(hermaphrodite) flowers. Due to this, the pollen grains have
more chances to come in contact with the stigma of the same
flower (This is called self pollination as discussed earlier).
Repeated and continued self-pollination results in
inbreeding depression (i.e., vigour and vitality of the plant
decreases).
To overcome this problem, flowering plants have developed
many devices to discourage self-pollination and to
encourage cross-pollination.
Some of the important devices are discussed below :
1. Dichogamy :This involves the maturation of male and
female organs at different time. In these cases, pollen release
and stigma receptivity do not occur at the same time. Either
the pollen is released before the stigma becomes receptive or
stigma becomes receptive much before the pollen release.
2. Herkogamy :Self-pollination is avoided by development
of some kind of physical barrier between stamen and pistil.
This may be achieved by placement of stamen and pistil at
some distance from each other in the same flower or by
inserting the anther within the corolla tube and extending the
style to the exterior of the flower, e.g., Gloriosa.
3. Heterostyly:It is a unique form of herkogamy in which
the lengths of stamen and pistil in a flower differ in
different plants of the same species, e.g., Primula.
4. Self-incompatibility: This is a genetically governed
mechanism to prevent self-pollination. In these cases, the
pollens of a flower are unable to germinate over the
stigma of the same flower.
5. Production of unisexual flowers :This is an important
mechanism to check self fertilisation. If both male and
female flowers are present on the same plant such as castor
and maize (monoecious), it prevents autogamy but not
geitonogamy. In many other species (as in papaya), male
and female flowers are present on different plants, that is
each plant is either male or female (dioecious). This
condition prevents both autogamy and geitonogamy.
Artificial Methods
Emasculation To carry out cross-pollination artificially, in a
bisexual plant (which does not show above adaptation), its
stamens are removed (cut off). This is called emasculation.
Emasculated flowers are covered with a suitable sized bags
(of butter paper) to prevent any contamination by unwanted
pollens. This process is called bagging.
Artificial Hybridisation
The process of artificial pollination, that leads to fertilisation
and formation of desired fruits is called artificial
hybridisation. It is one of the major approaches of crop
improvement programme.
Artificial Hybridisation in Bisexual Flowers
If the plant bears bisexual flowers, removal of anthers from
the flower bud before the anther dehisces using a pair of
forceps is necessary. This is called emasculation. To prevent
any contamination, bagging is also required. When the
flower attains receptivity, bags are removed off and mature
pollen grains (collected from anthers of the male parent) are
dusted on the stigma (artificial pollination). Now the flowers
are re-bagged, until the formation of fruit.
Artificial Hybridisation in Unisexual Flowers
If the plant produces unisexual flowers, there is no need for
emasculation. The female flower buds are bagged before the
flowers open (this is to avoid any unwanted pollination).
When the stigma becomes receptive, pollination is carried
out using the desired pollen and the flower is re-bagged as
above.
Pollen-pistil Interaction
ISC Biology-XII
Pollination does not ensure that the right type of pollen are
reaching the right type of stigma. It is purely a matter of
chance. Stigma can receive pollen of its own species or of
other species. But pollen of wrong type do not germinate on
the stigma. Only right type of pollen (.e., pollen of same
species) germinates on the stigma.
The pistil has the ability to recognise the type of pollen,
whether it is of the right type (compatible) or of the
wrong type (incompatible). If it is of the right type, the
pistil accepts the pollen and promotes post-pollination
events that lead to fertilisation.
If the pollen is of the wrong type, the pistil rejects the pollen
by preventing pollen germination on the stigma or the pollen
tube growth in the style.
The ability of the pistil to recognise the pollen depends upon
pollen-pistil interaction, which is genetically decided. This
interaction occurs with the help of certain chemical
substances.
Sexual Incompatibility
Stigma receives a variety of pollen grain but not all
pollen grains succeed to germinate and bring about
fertilisation. Stigma is so equipped as to allow only right
kind of mating types (pollen grains) to germinate over it.
The other pollen grains, which fail to germinate on a
stigma are said to be incompatible to it (stigma). This
phenomenon is known as 'sexual incompatibility'.
It can be of two types - Interspecific and intraspecific
incompatibility (or self-incompatibility).
1. Interspecific Incompatibility
In this type of incompatibility, the pollen grains and the
stigma, belong to two different species. It is thus heterogenic
incompatibility. It is controlled by more than one (different)
gene, located at different loci on chromosomes.
→ It prevents the cross-pollination.
In incompatible interspecific crosses, successful fertilisation
does not occur. If it occurs, it is followed by abortion of
hybrid embryo (due to inadequate development of
endosperm).
2. Intraspecific (Self) Incompatibility
In this type of incompatibility, the pollen grains and the
stigma belong to same species, i.e., the pollen grains of a
species fail to germinate on the stigma of the same species.
✦ In nature, different floral adaptations such as
dichogamy, herkogamy, unisexuality, etc. are found,
which prevent self-pollination.
→ But the most effective natural device is the self
incompatibility, which refers to the inability of the plant,
producing functional male and female gametes to let seeds,
when self pollinated.
If the incompatibility is due to the genotypes of the
sporophyte (or stigma tissues), it is known as sporophytic
incompatibility.
→ On the other hand, if the incompatibility is due to the
genotype of pollen (or gametophyte), it is known as
gametophytic incompatibility.
✦ It has been noted that the main cause of sexual
incompatibility is the gene, called S. It shows multiple
allelism. The various alleles are designated as S1, S2, S3
and so on.
(d) Development of Male Gametophyte after Pollination
It involves following steps:
(i) Formation of Male Gametes
(i) Only a right kind of spore germinates on the right kind of
stigma (i.e. for germination, both pollen and the stigma must
be of the plants, that belong to the same species).
(ii) After pollination, the male gametophyte (germinating
pollen) is usually at two cell stage. It contains a vegetative
cell and a generative cell.
(iii) The generative cell contains a large amount of
protoplasm. It acquires a characteristic vermiform
appearance. It now divides mitotically, to form two
elongated, haploid male gametes. (note: there is no meiosis,
because the dividing cell is already haploid).
(iv) Both these male gametes are genetically similar and
remain in pair for a considerable period of time. (ii)
Formation and Growth of Pollen Tube
(i) The exine (outer wall) of the pollen now ruptures and the
cytoplasm of the vegetative cell bulges out through germ
pore, in the form of a tube, called pollen tube.
(ii) Pollen tube at the apex contains tube nucleus (which is,
in fact, the nucleus of vegetative cell).
(iii) Behind the tube nucleus, there are 2 male gametes.
(iv) The tube grows towards the ovule (megasporangium),
making its way through the style.
(v) Usually, a single pollen tube arises from one pollen.
Such pollen grains are called monosiphonous. However
in some plants, pollen grains give rise to many pollen
tubes. Such pollens are called polysiphonous.
Now pollen tube makes its way through the style (of
carpel) and moves towards the mature ovule, containing
female gametophyte (embyo sac).
Haploid Vegetative
nucleus
cell
2 Male gametes (sperms)
Vegetative or tube nucleus
Pollen tube
Vegetative or tube nucleus
Microspore
Generative cell 2-celled
2 Male gametes (sperms)
3-celled, mature
Emergence of pollen tube
(pollen grain) male gametophyte male gametophyte
Fig. 2.16: Development of male gametophyte from a
microspore
(Note: Mature male gametophyte is 3-celled)
B. DEVELOPMENTS THAT LEAD TO THE
FORMATION OF
FEMALE GAMETE (EGG CELL)
The female reproductive whorl of the flower (i.e,
gynoecium) consists of megasporophylls (carpel or pistil).
Each pistil further consists of three parts:
Stigma
(a) Stigma: It is a terminal disc shaped portion, which
serves to receive pollens after their pollination.
Style
(b) Style: It is slender, hollow elongated tube, which
provides path to the pollen tube, for fertilisation.
(c) Ovary : It is a basal swollen part that internally contains
one or more ovules (or megasporangium).
Ovary
Fig. 2.17: Parts of pistil
Ovule internally produces megaspore (haploid cell), which
represents the first cell of the female gametophyte. The
megaspore on development produces mature female
gametophyte, (or embryo sac). This developmental process
can be studied in following steps:
1. Development of Ovule (Megasporangium)
Ovules or megasporangia are developed within the cavity of
ovary (locule) and remain attached through a placenta with
the ovarian wall. Ovule arises as a small mound of
homogeneous tissue on the inner wall of the ovary. This
mound develops to form the inner central part of the ovule,
called nucellus. This mound becomes covered with two
integumentary processes.
Structure of Ovule (Megasporangium)
Antipodals
Chalaza pole
(i) Each ovule is attached to the inner wall of the ovary
(placenta), by a slender stalk, called funicle. (ii) The point of
attachment of ovule to its funicle is calledhilum. (iii) Main
body of the ovule is formed by inner central mass i.e., Inner
nucellus. Nucellus consists of living parenchymatous cells.
(iv) In mature ovules, the nucellus serves to cover and
provide nutrition to the embryo sac (female gametophyte).
- (iii) The parietal cell may either remain undivided or
(v) Each ovule has two distinct ends (a) micropyle end (it is
also called opening of ovule. It is the end wherefrom the
pollen tube usually enters the ovule during fertilisation) and
(b). chalaza end (the posterior end, opposite to micropylar
end)... (vi) Externally the nucellus is covered by one or two
protective
-Outer integument
Embryo sac
Synergids
integument
Secondary nucleus Nucellus
Egg
- Hilum
-Funicle
Micropyle
Fig. 2.19: Structure of ovule
covers, called integuments. These integuments arise from
the chalazal end. (vii) When only one integument is
present, the ovule is called unitegmic, and if the ovule
consists of two integuments, it is calledbitegmic. In some
plants such as Santalum, Olax etc,ategmic (no
integument) condition may be present.
(viii) In mature ovules, the female gametophyte or embryo
sac is present in the centre. The embryo sac consists of egg
cell (female gamete), synergid cells, antipodal cells and
polar nuclei (This is described a little later).
Types of Ovules (Not in syllabus)
On the basis of position of micropyle, with respect to the
funiculus, ovules are of 6 types:
1. Orthrotropous Ovule
It is atropous or straight, where the micropyle, chalaza and
the funiculus, all are in the same line, e.g., Cycas, Family
Polygonaceae and Piperaceae.
2. Anatropous Ovule
It is of the most common occurrence. In this ovule, the
funicle is long and whole body of the ovule is inverted,
through 180°. As a result the micropyle comes close to
the funicle, e.g., Most common in dicots and monocots. 3.
Hemianatropous or Hemitropous Ovule
In this case the body of the ovule is inverted only through
90°. As a result the funicle comes to lie at right angle to the
nucellus. Micropyle and chalaza, lie in the same plane, e.g.,
Ranunculus.
4. Campylotropous Ovule
When body of the ovule is not completely inverted, but it is
bent like 'horse shoe'. The micropyle and chalaza do not lie
in the same plane (however the nucellus/embryo-sac remain
straight), e.g., Family Capparidaceae, Cruciferae etc.
5. Amphitropous Ovule
It is similar to campylotropous, but in this case the
nucellus/embryo sac is also bent like 'horse shoe', e.g.,.
Family Alismaceae.
6. Circinotropous Ovule
It is of a very rare occurrence. Here the body of the
ovule is bent through 360°, so that it takes a one
complete turn. (Micropyle, chalaza and the nucellus are
all in same plane), e.g., Opuntia.
2. Formation of Megaspore (Megasporogenesis)
Circinotropous
(i) In the micropylar region of the nucellus, usually a single
cell gets differentiated from other cells. It is called primary
archesporial cell. It contains a big nucleus, dense cytoplasm
and has a larger size. (ii) This cell divides periclinally, to
form primary parietal cell and primary sporogenous cell.
undergoes a few periclinal and anticlinal divisions, so that
the sporogenous cell gets embedded in the nucellar mass.
Consequently, the sporogenous cell becomes sub-
hypodermal in position. Ovules with such a hypodermal
sporogenous cell are called
crassinucellate.
(iv) The sporogenous cell now acts as megaspore mother
cell (MMC). It undergoes reductional division (meiosis)
and forms four haploid megaspores. This process is
called megasporogenesis. (v) In most of the angiosperms,
out of these 4 megaspores, 3 get degenerated (to provide
more nourishment to the remaining one). Thus, only one
megaspore remains functional in the ovule. This gives
rise to female gametophyte on development. Such female
gametophyte (embryo sac) which develops from a single
megaspore, is called monosporic embryo sac. This
development is called monosporic development (In some
angiosperms bisporic or even tetrasporic embryo sac
may also be present). (vi) A megaspore is a haploid
structure and represents the first cell of the female
gametophyte. It develops to form fully matured
gametophyte. (Megaspore should not be considered as a
female gamete).
Megaspore mother cells
Nucellus
M1
Meiosis
M2
4 Megaspores
3 spores degenerate
Functional megaspore
Fig. 2.21: Formation of megaspore (Megasporogenesis)
3. Formation of Female Gametophyte (Embryo sac)
Female gametophyte is also called embryo sac. It develops
from megaspore. There are great variations in the
development of embryo sac. Some are monosporic, some
others are bisporic and rest others are tetrasporic, as
discussed above (We are describing here the development of
a typical embryo sac, which is monosporic and is of the
most common occurrence in angiosperms).
(i) The remaining megaspore in the ovule is given sufficient
nourishment, so that it becomes larger in size. (ii) This
megaspore now divides by three successive mitotic divisions
and forms 8 nuclei. All these nuclei are haploid. These
mitotic divisions are strictly free nuclear, that is, nuclear
divisions are not followed immediately by cell wall
formation.
Haploid megaspore
→ 2 nuclei
(iii) Out of these 8 nuclei, 3 nuclei (at micropylar end),
undergo cytokinesis (forming cell wall) and form egg
apparatus. (iv) The egg apparatus contains 2 synergid
cells and one egg
cell.
4 nuclei
8 nuclei
Egg apparatus
The three cells at micropylar end, i.e., egg cell along with
two synergids, together, are called egg apparatus.
(v) The egg cell represents the female gamete.
(vi) The other three nuclei (at chalazal end) also undergo
cytokinesis and form three antipodal cells.
(vii) Remaining two nuclei are present in the centre (they do
not follow cytokinesis). These are called central or polar
nuclei.
(viii) This entire structure is called embryo sac, which
represents the mature female gametophyte. Thus,
normally it contains 3+2+3 arrangement of nuclei in a
typical embryo sac.
ISC Biology-XII
3 Antipodals
2 polar nuclei
Synergids
Egg cell (female gamete)
(ix) The mature female gametophyte (embryo sac)
consists of 8 nuclei, but only 7 cells (one egg cell, 2
synergids, 3 antipodal cell and one Fig. 2.22: Mature
embryo sac remaining cell, in which 2 polar nuclei are
present).
Structure of Mature Embryo sac (Female Gametophyte)
As said earlier, the mature female gametophyte or embryo
sac in a typical angiosperm is 7 celled and 8 nucleate. It
contains one egg cell, 2 synergids, 3 antipodal cells and 1
largest central cell with 2 polar nuclei. synergids
These are 2 elongated cells, present at the micropylar end of
the ovule, one on each side of the ovule. The characteristic
feature of the synergid cells is the presence of a filiform
apparatus.
Filiform apparatus
It is a mass of finger like projections, which are present in
the synergid cells. These projections are the in growths of
the cell wall. These are rich in polysaccharides. Each
projection is provided with a set of microfibrils. It is
supposed to be developed due to some chemotactic stimulus.
It helps in the short distance transport of nutrients across the
synergid wall.
(i) Synergids play an important role in directing the pollen
tube growth, by secreting some chemotropically active
substances.
(ii) The degenerating synergids help pollen tube to discharge
and release its contents, in the embryo sac. Some workers
have suggested its haustorial or nutritive function.
Antipodals
(i) These are usually 3 in number, present at the chalazal end
of the embryo sac.
(ii) These exhibit the great variation in size, structure, life
span and biochemistry.
(iii) Usually, these cells degenerate before/soon after
fertilisation (But in Caltha pulustris the antipodals persist
upto the octant stage of the pro-embryo).
(iv) Antipodal cells serve to provide nutrition to the pro-
embryo. Egg Cell
(i) It is a single haploid cell, lies at the micropylar end,
between two synergids.
(ii) It represents the female gamete.
Chalazal end
-3 Antipodal cells
Micropylar end
2 polar nuclei
Central cell
-1 Egg cell
Egg
apparatus
2 Synergids
Filiform apparatus
Fig. 2.23: A typical embryo sac in angiosperms
(iii) It when fertilised by sperm (male gamete), forms
diploid zygote.
(Female gametophyte)
Table 2.4: Differences between Male and Female
Gametophytes in Angiosperm
S.No.
Male Gametophyte
It is derived from a pollen grain or microspore.
Female Gametophyte
It is derived from a megaspore.
41
2
3.
5.
6.
7.
It does not remain permanently embedded inside the The
female gametophyte remains permanently microsporangium.
embedded in the megasporangium or nucellus.
The male gametophyte comes out of the pollen grain by The
female gametophyte remains surrounded by the forming a
pollen tube.
The male gametophyte is only 3-celled.
All the cells of the male gametophyte are functional. The
tube cell is required to carry the two male gametes, both of
which take part in fertilization.
membrane of the megaspore.
The female gametophyte is 7-celled.
The antipodal cells do not seem to perform any function
except absorption of nourishment from nucellus in certain
cases. Out of two synergids only one is required for
receiving the pollen tube.
The remains of male gametophyte disintegrate after After
fertilization two new structures are produced both
fertilization. of which show active growth.
The gamete produced by male gametophytes are The gamete
produced by female gametophyte is egg
sperms.
cell.
C. FERTILISATION
Fertilisation in angiosperms, is unique. It is called double
fertilisation. This is because there are two sperms per pollen
tube. One of the sperms fertilises the egg cell to form
zygote, while remaining sperm fertilises two polar nuclei,
resulting in the formation of a triploid endosperm nucleus.
Important events in fertilisation are following:
1. Entry of Pollen Tube into Ovule and Embryo sac
(i) After arriving in the ovary, the pollen tube finds its way
through style and enters the ovule.
(ii) In majority of the cases, the pollen tube, enters the
ovule through the micropyle and then enters one of the
synergids through the filiform apparatus. The entry of
pollen tube through micropyle is called porogamy.
(iii) Filiform apparatus present at the micropylar part of the
synergids guides the entry of pollen tube.
(iv) All these events from pollen deposition on the stigma
until pollen tubes enter the ovule, are together referred
to as pollen-pistil interaction.
Pollen grains
Signa
Style
Polen tube
Embryo sac
-Antipodal
Polar nudei
Egg cell
Synergid
Micropyle
Ovule
Fig. 2.24: Longitudinal section of a flower showing growth
of pollen tube
(v) In some plants, the pollen tube enters the ovule through
chalazal end, however the entry into the embryo sac occurs
through micropyle, e.g., Casuarina. This is called
chalazogamy. In a few plants, the pollen tube enters the
ovule through the funiculus or integuments. This is called
mesogamy. However the final entry into the embryo sac
occurs through micropyle.
2. Discharge of Male Gametes
(i) The pollen tube contains two (haploid male gamete).
(ii) When pollen tube enters the embryo sac (inside the
ovule), it bursts to release its contents i.e., two sperms
along with certain amount of protoplasm.
(iii) Now the sperms are ready for fertilisation.
(iv) In angiosperms, the fertilization involves (a) syngamy
and (b) triple fusion. Hence it is also called as
double fertilisation.
3. Double Fertilization
Double fertilization is the characteristic feature of
angiosperms. It was first observed by Nawaschin (1898) in
Fritilaria and Lilium. It involves two types of fusion (a)
syngamy (fusion of egg cell and one sperm)
and (b) triple fusion (fusion of remaining sperm and two
polar nuclei).
(1) Syngamy
One of the two sperms goes to fertilise the egg cell. This
fusion is called syngamy. It results in the formation of
zygote, which gives rise to proper embryo.
(ii) Triple fusion
The remaining sperm now fuses with the two haploid polar
nuclei (present in the centre of embryo sac). This fusion is
called as triple fusion (as three nuclei i.e., one sperm and 2
polar nuclei, are fused). It results in the formation of triploid
Primary endosperm nucleus (PEN), which on development
gives rise to Primary endospermal cell (PEC) and then
endosperm. Endosperm is therefore, triploid in angiosperms
(It is a characteristic feature of angiosperms). Endosperm
serves to provide nutrition to the developing embryo.
1 Sperm + Egg cell 1 Sperm + 2 polar nuclei
Zygote (2x)
= Syngamy
Double fertilisation
Pri. Endosperm nucleus (3x)
= Triple fusion
Sperms
Polar nuclei
Egg cell
Pollen tube bursts and releases 2 sperms
Sperms
Triple fusion
(2 fused polar nuclei +
Sperm)
Double fertilisation
Antipodal and synergids degenerate
Primary endosperm
nucleus (3x)
Syngamy (Egg cell + Sperm)
Zygote (2x)
Fig. 2.25: Release of sperms and double fertilisation
Significance of Double Fertilization
Double fertilisation is necessary for the formation of
viable seeds, as it ensures the formation of endosperm,
together with the embryo.
✦ Endosperm (which is formed by the triple fusion) is
essential for the proper growth of the embryo, as it
(endosperm) provides nourishment to the embryo.
D. POST-FERTILISATION STRUCTURES AND
EVENTS
45
After double fertilisation, the ovule undergoes further
development. Internally there is endosperm formation,
development of the embryo (sporophyte). The ovule
gradually becomes the seed and ovary becomes the fruit.
All these events are collectively known as post-
fertilisation events.
ENDOSPERM
Endosperm development occurs before the development of
the proper embryo, this is because the development of the
embryo requires heavy nutrition, which is supplied by the
endosperm. So, endosperm is a sterile but nutritive tissues.
After fertilisation, the primary endosperm cell (containing
PEN) divides repeatedly and forms a triploid endosperm
tissue. The cells of endosperm contains large amount of
reserve food, which is utilized by the cells of embryo for
their development.
Endosperm depending upon the mode of its formation,
endosperm in angiosperms is of three types-nuclear, cellular
and helobial.
1. Nuclear endosperm: it is most common type of
endosperm. The primary endosperm nucleus divides
repeatedly without wall formation to produce a large
number of free nuclei. Now cell walls are laid by
simultaneous cytokinesis to form a multicellular endosperm.
This type of endosperm is found in wheat, rice, maize,
sunflower etc.
2. Cellular endosperm: Every division of the primary
endosperm nucleus is followed by cytokinesis. Thus, cell
walls are laid successively. In this way, endosperm remains
cellular right from the beginning. This type of endosperm is
found in Datura, Petunia and Balsam.
3. Helobial endosperm: It is of intermediate type between
cellular and nuclear endosperms. The first division of
primary endosperm nucleus is followed by transverse
cytokinesis to form two unequal cells, one larger cell
towards micropylar end and one smaller cell towards
chalazal end. Further development of both the cells occurs
like that of nuclear endosperm. The chalazal chamber often
remains smaller and may degenerate. This type of
endosperm is found in order helobiales of monocots.
Points to remember
Primary
endosperm
nucleus
(a) Nuclear endosperm
Free nuclei
Primary endosperm nucleus
(b) Cellular endosperm
Primary endosperm nucleus
(c) Helobial endosperm
-Micropylar chamber
-Free nuclei
Endosperm
Chalazal chamber
Cell wall formation
Endosperm
Fig. 2.27 (a): Types of endosperm
(i) The endosperm in angiosperms is the result of triple
fusion (sperm + 2 polar nuclei) and has 3x nature (triploid).
Since it is triploid, it is sterile in nature. Triploid nature of
endosperm makes it genetically useless.
(ii) The endosperm provides the essential food to the
developing embryo.
(iii) Thus the highly nutritive nature of endosperm in
angiosperms makes them highly successful plants as they
can provide a good deal of nutrition to their embryo.
(iv) In some families of angiosperms (order - Helobiales), a
special type of endosperm is found. This type of endosperm
development is intermediate between nuclear and cellular
type of endosperm development. It is called helobial type of
endosperm.
EMBRYO
After double fertilization, each ovule develops into a seed.
Seed is therefore a ripend ovule. It contains an embryo.
Embryo develops from the zygote, so it is diploid
(sporophytic) in nature. Embryo is developed at the
micropylar end, where the zygote is situated.
The development of the zygote into an embryo takes place
only after certain amount of endosperm is formed. This is an
adaptation to provide assured nutrition to the developing
embryo.
The early stages of embryo development (embryogeny) are
similar in both monocotyledons and Dicotyledons, however
their seed structure differs greatly.
The zygote gives rise to the proembryo and subsequently to
the globular, heart-shaped and mature
embryo.
Development of embryo: Dicot embryogenesis
- embryonal axis and two cotyledons.
(i) Zygote divides by a transverse division, forming two
cells (i) Basal cell and (ii) Terminal cell. (ii) The basal cell
divides by transverse division to form many cells, arranged
in a row. This structure is called suspensor. The function of
suspensor is to push the proembryo into the embryo sac
cavity and to absorb and transfer nutrients to the proembryo.
(iii) The lower most cell of the row is differentiated as
hypophysis, which gives rise to radicle. (iv) The terminal
cell divides by 2
vertical and one transverse divisions to form eight cells.
These are arranged in two tiers of 4 cells each.
(v) Upper tier (just below the suspensor) gives rise to
hypocotyl. The lower tier gives rise to epicotyl, cotyledons
and plumule.
(vi) This entire structure is called pro-embryo.
Now,
this
structure grows further and becomes heart shaped.
Zygote
Basal cell
Haustorial cell
Suspensor
(a)
Terminal cell
Hypophysis
(b)
(c)
Tetrad
Octant
(d)
(e)
Reduced suspensor
Mature embryo
Suspensor
(vii) In dicots, two cotyledons are formed, but in monocots
only
Hypocotyl
Epicotyl
Cotyledons
one cotyledon develops.
(viii) By the time, the integuments of the ovule, become
hard and are called as seed coat and the
entire ovule is now called as seed.
(ix) When seed germinates, the
Pro-embryo (1)
Heart-shaped stage (g)
(h)
Cotyledons
Fig. 2.28: Various stages of the development of embryo
(in dicots)
cotyledons are used up in providing nourishment. The
radicle gives rise to root system, while plumule gives rise
to shoot system.
Basic structure of embryo
An embryo consists of an embryonal axis. In the middle
of the embryonal axis, cotyledons are present (one in
monocots and two in dicots). The part of embryonal axis,
above the level of cotyledons is called epicotyl, whereas
the part, below the level of the cotyledons is called
hypocotyl. The tip of epicotyl is called plumule, which on
development gives rise to shoot of the plant. Hypocotyl
ends into radicle, which on development, gives rise to
roots.
A typical dicotyledonous embryo
(1) A typical dicotyledonous embryo consists of an
(ii) The portion of embryonal axis above the level of
cotyledons is the epicotyl, which terminates into plumule or
stem tip. It gives rise to shoot of the plant and it is
negatively geotropic.
(iii) The cylindrical portion below the level of cotyledons is
hypocotyl that terminates at its lower end in the radicle or
root tip. It gives rise to root of the plant and it is positively
geotropic.
(iv) The root tip is covered with a root cap.
Development of embryo in monocots: Monocot
embryogenesis (in corn, Zea mays)
(i) Monocots have a more complex embryo structure in the
mature seed compared to dicots, but early embryonic
development is similar to that of dicots.
(ii) In the zygote, the first cell
division is transverse and asymmetrical. It leads to
formation of an apical cell and a basal cell.
(iii) The basal cell is larger and lies towards the micropylar
end. It does not divide divide again but becomes
transformed directly into a large vesicular cell.
(iv) The
apical cell divides transversely forming two cells, of these,
the lower cell divides vertically forming two cells. These
cells once again divide vertically forming a group of four
cells (quadrant stage).
Basal cell
Apical cell
Zygote
F
Vesicular cell
60
E
R
L
Vesicular cell 4.
Suspensor Style and stigma
R 5.
R Ovary
Radicle 6.
Cotyledon Ovary wall
H 7.
Fig. 2.30: Stages in the development of a typical monocot Ovule
embryo in Sagittaria 8.
(v) The cell next to the quadrants also divides vertically and Funicle
the cell next to the upper vesicular cell divides several times 9.
transversely. Hilum
(vi) The quadrants now divide transversely giving rise to an 10.
eight-celled embryo (octant stage), in which the cells are Nucellus
arranged in two tiers of 4 cells each. 11.
(vii) Each cell of the octant stage divide periclinally, giving Outer integument
rise to the dermatogen. 12.
(viii) Later the periblem and plerome are also differentiated. Inner integument
All these regions, derived from the octant stage, later give 13.
rise to a single terminal cotyledon. Micropyle
A typical monocotyledonous embryo 14.
(i) Embryos of monocots possess only one It is generally Synergids
called 15.
cotyledon. scutellum. Egg cell
(ii) The cotyledon is situated towards one side (lateral) of 16.
the embryonal axis. (iii) At its lower end, the embryonal Antipodal cells
axis has the radicle and root cap. 17.
(iv) The radicle and root cap remain packed in a protective Secondary nucleus
undifferentiated sheath (cover) called coleorhiza. SEED
(v) The portion of the embryonal axis above the level of Seed can be defined as a mature ovule. It is the final product
attachment of cotyledon (scutellum) is called epicotyl. of sexual reproduction in an angiospermic plant.
(vi) Epicotyl has a a Structure of a Typical Dicot Seed (Bean Seed)
plumule. (i) The bean seed is kidney shape. (ii) The outer seed coat,
coleoptile. called testa is smooth and brown or red. The inner seed coat,
shoot apex, called called tegmen, is thin and white. It is very difficult to
2 Cotyledons separate the testa from the tegmen.
(a) (iii) An oval scar on the concave side marks the hilum. It
Shoot apex (plumule) represents the place where the seed was attached to the
Embryonal axis fruit-wall. Near one end of the hilum is a tiny pore, the
Radicle micropyle.
Root cap Fall off
Scutellum (cotyledon) Fall off
Coleoptile Fall after dehiscence of anthers
Shoot apex (plumule) Fall off
Epiblast Formation of fruit
Embryonal axis Pericarp
Radicle Seed
Root cap Stalk of the seed
Coleorhiza Hilum
(b) Perisperm
Fig. 2.31: (a) Dicot embryo, (b) Monocot embryo Testa
(vii) The plumule bears a few leaf primordia (early leaves) Tegmen
enclosed in a hollow foliar structure, called Micropyle
Table 2.5: Fate of different floral parts after fertilization Degenerate
(Changes in the ovary and ovule for fruit and seed Embryo
formation) Degenerate
Before fertilization Endosperm
After fertilization Seed coat
S.No. Apical meristem Epicotyl
1. First leaves
Sepals -Hypocotyl
234 Embryonic root
2. Apical meristem
Petals Cotyledons
3. Embryo
Stamens Fig. 2.32 Structure of a typical dicot (Bean) seed
(iv) Below the seed coat, two thick and fleshy cotyledons
are present. The cotyledons contain stored food in the form
of starch, protein and oil.
(v) The embryo is present between the cotyledons. Its
narrow end is called the radicle which gives rise to the root
on germination. The small feathery structure at the opposite
end is the plumule which gives rise to
the shoot.
(vi) The region situated below the point of attachment of the
cotyledons, is called hypocotyl, and the region above it is
called the epicotyl.
Structure of a Typical Monocot Seed (Maize Seed)
Each maize grain is actually a one-seeded fruit in which the
testa is inseparably fused with the fruit wall (pericarp).
The outermost layer is formed by the fusion of the fruit-wall
and seed-coat. In a longitudinal section, the upper one-third
region constitutes the endosperm. It stores reserve food in
the form of starch. The endosperm is surrounded by a thin
aleurone layer. The aleurone layer stores proteins and fats.
the
endosperm
The area below represents the single cotyledon. It is known
Endosperm
Embryo
External view
Seed coat
& fruit-wall Aleurone - layer
49
Endosperm
Scutellum
Coleoptile
Plumule
Radicle Coleorhiza
Longitudinal section
Fig. 2.33: Structure of a monocotyledonous seed (Maize)
as the scutellum. It is separated from the endosperm by
the epithelial layer. The embryo is embedded in the
scutellum. Its plumule is covered by a sheath called the
coleoptile and the radicle by the coleorhiza.
Endospermic/Non-endospermic Seed
(1) Endospermic (or albuminous) seeds: The endosperm is
present in the mature seed and serves as food storage organ.
Examples: Castor, maize, onion.
(ii) Non-endospermic (exalbuminous) seeds: The cotyledons
serve as the only food storage organs. During embryo
development the cotyledons absorb the food reserves from
the endosperm. The endosperm is almost absent in the
mature seed. Examples: gram, pea, mustard.
Perisperm
In seeds of some plants, the nucellus persists after
fertilization. This persistent nucellus is called perisperm. It
provides nutrition to the seed. Examples: Castor, cotton,
beet, black pepper, coffee, etc.
Function of seed (Significance of seed formation)
Seed serves two important purposes -
(i) Dispersal of the species to new locations (aided in
angiosperms by the fruit). It avoids over-crowding and intra-
specific competition.
(1) Survival of the species during unfavorable dimatic
conditions (winter and excessive summer). Annual plants
such as beans, cereal grains, many weeds etc., can survive
through colder periods, only as seeds. When conditions
become favorable, germination occurs and a new generation
of plants develops.
Seed Dormancy
Most of the seeds do not germinate soon after their
formation. This period during which they do not germinate,
is called dormancy period. This phenomenon is called seed
dormancy. This is mainly due to, hard seed coats
(integuments), lack of water contents, lack of growth
promoting hormones, small size of micropyle due to which
oxygen availability is poor. In most of the seeds, dormancy
can be broken by providing optimum conditions for
germination.
Dormancy is beneficial for the plant in two ways:
(a) Dormancy is a mechanism to prevent germination
during unsuitable ecological conditions, when the
probability of seedling survival is low.
(b) One important function of dormancy is to delay
germination process, which allows enough time for
dispersal and spread of seeds to different areas.
Dormancy and Seed Viability
ISC Biology-XII
During dormancy period, the immature embryo remains
alive and seed remains viable. However, the period of this
viability varies greatly in different plants. Some seeds loose
their viability in just few days or months, while other
remains viable for many years. We have several records of
very old but viable seeds. The oldest record is that of a
lupine plant (Lupinus arcticus). The seeds were excavated
from Arctic Tundra and they germinated and flowered after
an estimated time of 10,000 years of dormancy. One more
notable record is that of date palm (Phoenix dactylifera),
showing about 2000 years of dormancy.
Some Important Terms, Related to Seeds and Fruits
Carpophore-Floral axis extension between adjacent
carpels, as in the Apiaceae. Ectocarp or Exocarp -
Outermost layer of pericarp.
Endocarp-Innermost differentiated layer of pericarp.
Funiculus - Seed stalk.
Mericarp - A portion of fruit that seemingly matured as a
separate fruit. Mesocarp - Middle layer of pericarp.
Pericarp - Fruit wall.
Placenta Thalamus
-
-
Region of attachment of seeds on inner fruit wall.
Base of the flower.
Replum - Persistent septum after dehiscence of fruits, as in
the Brassicaceae.
Retinaculum, Jaculator or Echma - A persistent indurated,
hook-like funiculus in the fruits of family Acanthaceae.
Rostellum or Beak - Persistent stylar base on fruit.
Seed - A matured ovule.
Septum or Dissepiment - A partition.
FRUITS
Fruits are formed as a result of development of the ovary
wall and other flower parts. In simple words, a fruit is a
ripened (mature) ovary. As seeds mature, they release the
hormones (mainly cytokinin), which stimulates the wall of
the ovary to develop into the fruit.
Development of Fruit
After an ovule is fertilized, the ovary begins to expand. The
petals of the flower, fall off and the ovule develops into a
seed.
The wall of the ovary surrounding the seed develops rapidly,
under the effect of hormones like cytokinin and becomes
enlarged in size. Fruit development continues until the seeds
have matured. The extent of development of the flesh of the
fruit is proportional to the number of fertilized ovules.
The wall of the fruit, developed from the ovary wall of
the flower, is called the pericarp. The pericarp is
differentiated into three distinct layers :
(a) Exocarp (the outermost layer - also called epicarp),
(b) Mesocarp (middle layer), and
(c) Endocarp (inner layer).
Some fruits, especially simple fruits, are derived from an
inferior ovary, so that other parts of the flower (such as
petals, sepals, and stamens), fuse with the ovary and ripe
with it. When such floral parts are a significant part of the
fruit, it is called false fruit.
So, a false fruit is that which do not develop from the ovary
or in which the significant part is derived from floral parts,
other than ovary, e.g., Apple, cashewnut.
In apple and pear, thalamus is modified to form fruit. In
strawberry, the receptacles become fleshy and forms the
main part of the fruit.
True and False Fruits
If the fruit is developed from the ovary, it is called true fruit.
But when it develops from any other floral parts (such as
thalamus, receptacle or even calyx) then such a fruit is
called a false fruit, e.g., in apple and pear, thalamus is
modified to form fruit. The structures actually derived from
the ovary, are not edible. Similarly in cashewnut, the
peduncle is modified to form fruit.
Seed
Thalamus
Fig. 2.34:False fruit of apple (edible part consists of
thalamus)
FRUITS
True fruits (Develop from ovary
of a flower)
Simple fruits (Develop from
single ovary
of single flower)
Dry fruits
(Pericarp is dried,
e.g., Guava, orange,
Fleshy fruits
False fruits
(Develop from floral parts,
other than ovary, e.g., Apple strawberry, Cashewnut)
Aggregate fruits or etaerio (Develop from multicarpellary
gynoecium, where
carpels are fused together), e.g., Raspberry
(Pericarp is fleshy, moist or fibrous),
e.g., Groundnut, mustard
Fig. 2.35: Classification of various types of fruits
Parthenocarpic Fruits
Parthenocarpic fruits (Develop without fertilization), e.g.,
Banana
Multiple fruits (Develop from flowers of whole
inflorescence), e.g., Mulberry
(Development of fruit without fertilisation is called
parthenocarpy and such fruits are called parthenocarpic
fruits. The fruit resembles a normally produced fruit but is
seedless i.e., a parthenocarpic fruit does not have an embryo
and endosperm. Banana is an important example of such
fruits.
Certain varieties of the pineapple, cucumber, grape, orange,
grapefruit, persimmon and breadfruit are also good
examples of naturally occurring parthenocarpic fruits.
Parthenocarpy can be induced in non-parthenocarpic
varieties also. Several edible fruits, such as seedless grapes,
oranges etc., have been developed to afford the taste of the
fruit without the seed. Parthenocarpic fruit do not have
seeds, so they can not be used to raise viable offspring.
Structure of Some Fruits
Mango and coconut belong to the category of drupe fruits.
1. Mango
Exocarp
Mango is a common example of a simple fleshy drupe fruit.
It is the national fruit of India. Let us consider the structure
of a mango fruit.
-Mesocarp
-Endocarp
(i) Epicarp: It is thin and leathery.
(ii) Mesocarp: When the epicarp is peeled off, the thick and
fleshy mesocarp is exposed. This is the edible part of mango
fruit.
(iii) Endocarp: It is hard and stony. It forms a covering
around the seed. (iv) Seed: Mango has a single seed, which
is elongated and laterally compressed.
Fig. 2.36:Fleshy drupe of mango
2. Coconut
Coconut is a type of fibrous drupe.
(i) Epicarp: It is thin but tough.
(ii) Mesocarp: It is thick and fibrous.
(iii) Endocarp: It is thick and woody. This layer forms a
covering around the seed.
(iv) Seed: The seed is large in size. It is composed of a very
thin testa, lined by a thick layer of white endosperm which
is edible.
Dispersal of Fruits and Seeds
Shoot- (stem & leaves) Mesocarp (fibrous husk) Root-
ISC Biology-XII
Coconut "apple
A sweet, spongy mass (cotyledon) that dissolves &
absorbs the endosperm Coconut "meat" (endosperm)
Endocarp
(Surrounding seed)
Exocarp
(outer layer)
Mesocarp
(fibrous husk)
Fig. 2.37: Fibrous drupe of coconut
Plants are fixed to a substratum and are unable to move. All
their life activities such as growth, production of flowers
and fruit etc, occur at the same place. Consequently, the
fruits and seeds usually fall directly under the mother plant.
Such seeds and fruits have to germinate and develop under
limited food supply and space. But to solve this problem,
there are many devices to disperse fruits and seeds to distant
areas. Some important devices are wind, water and animals
(insects and birds etc.).
Enormous reproductive capacity of flowering plants
Flowering plants reproduce at a very high rate in order to
disperse and spread their species. Some, plants have
enormous reproductive capacity. This is achieved by
producing large number of flowers, ovules, embryo sacs and
seeds.
Orchid fruits contain thousands of tiny seeds. Similarly
some parasitic species such as Orobanche and Striga also
produce large number of tiny seeds. A single Ficus tree can
produce billions of seed.
SPECIAL MODES OF REPRODUCTION
(Reproduction without normal fertilisation)
In the above text we discussed the normal reproduction
pattern in a typical angiosperm. But there are many
cases where new plants can be produced in an unusual
manner. Three important methods by which new plants
can be produced unusually are - Apomixis, feature is only the presence of more than one embryo, per
polyembryony and parthenogenesis. ovule (seed)].
Apomixis Polyembryony is very common in conifers. Many dicot
Vegetative reproduction (e.g., orange, lemon, jamfolan, mango, groundnut) and
Agamospermy monocot (e.g., onion) species also exhibit this feature.
Unusual modes of reproduction Polyembryony may be due to following reasons:
Polyembryony (i) Presence of more than one egg cells in the embryo sac.
Tissue culture (ii)(Synergids or antipodals start acting as egg cell.)
methods (iii) Normal zygote may divide into too many fragments,
Diplospory each acting as a fresh zygote.
Parthenogenesis (iv) Presence of more than one embryo sac in an ovule.
Apospory Polyembryony may also be induced in normal plants by
Adventive embryony artificial methods, e.g., treatment of freshly harvested seeds
Fig. 2.38: Classification of various unusual modes of with chemical like 2,4 dichloro phenoxy acetic acid may
reproduction induce polyembryony.
1. Apomixis
(Formation of new plants without fertilisation)
Almost all plants show clear alternation of generation from
gametophyte (haploid) to sporophyte (2x) and vice-versa. In
angiosperms, the sporophytic (2x) stage is highly dominant,
while the gametophytic stage is very short. Sometimes, the
deviation may occur in the life cycle of a plant from clear
alternation of generation. This deviation is referred to as
apomixis.
(Apomixis may be defined as 'the substitution of normal
sexual reproduction by another form of reproduction, that
does not involve meiosis and syngamy (fertilisation). Plants
showing this phenomenon are called apomictic) 2.
Parthenogenesis
(The direct formation of embryo from female gamete (egg)
without any fertilisation is known as parthenogenesis.)
Parthenogenetic products may be haploid, diploid or
polyploid. Parthenogenesis involves following subtypes :
(1) Diplospory: In this case, the megaspore mother cell (2x)
does not undergo meiosis and directly starts behaving as
megaspore. Thus, megaspore is diploid and it gives rise to
diploid embryo sac (female gametophyte). The diploid egg
cell of embryo sac starts germination (without fertilisation)
as zygote and gives rise to diploid embryo.
Parthenocarpy
It refers to the development of fruit in an unfertilised flower.
It results in a seedless fruit. It may occur naturally (as in
banana, some varieties of apple, pear, pineapple etc) or may
be induced artificially by the application of hormones (as in
tomato).
(ii) Apospory: It involves the development of diploid
embryo sac from a normal diploid cell of plant (somatic cell,
other than megaspore mother cell). These sporophytic
tissues may be of nucellus or of integument etc. The diploid
egg cell of embryo sac starts germination (without
fertilisation) as zygote and gives rise to diploid embryo.
In both these cases, the embryo sac (of female gametophyte
is diploid, which is an unusual feature). Adventive
embryony : In this case the embryo develops inside the
ovule from any tissue (2x) directly, but not from the embryo
sac. Thus in other words, sporophytic generation directly
gives rise to new sporophytic generation. The sporophytic
tissue forms a bud like structure that directly starts behaving
as an embryo. (This is called sporophytic budding). Embryo
sac (female gametophyte) does not form in this condition.
3. Polyembryony
(The occurrence of more than one embryo in a seed is called
polyembryony) It was discovered by Leeuwenhoek (1917).
[Students should note that in polyembryony, normal gamete
formation and fertilisation are involved, but the unusual