1

II PU/CLASS XII

CHAPTER 2: SEXUAL REPRODUCTION IN FLOWERING PLANTS

ANGIOSPERMS are flowering plants. Flowers are the parts which facilitate their
reproduction. Flowers develop after completion of vegetative phase.
Flowers are the reproductive organs of the plant.
PARTS OF A FLOWER:-
A typical flower is made up of 4 whorls – calyx, corolla, androecium, and gynoecium.
These 4 whorls are arranged on the thalamus.
Calyx: the outermost and the first accessory whorl of a flower. Individual units are called
sepals.
Corolla: the second accessory whorl of a flower. Individual units are called petals.
Androecium: The first essential whorl. It represents the male reproductive part of a
flower. Androecium is made up of stamens.
Gynoecium (pistil) is the female reproductive part made up of carpels.

PRE-FERTILISATION: STRUCTURES AND EVENTS

Several hormonal and structural changes are initiated which lead to the differentiation
and further development of the floral primordium. Inflorescences are formed which bear
the floral buds and then the flowers.
• In the flower, the male and female reproductive structures, the androecium and
the gynoecium differentiate and develop.

ANUSREE M NAIR
 2
II PU/CLASS XII

• Androecium consists of a whorl of stamens representing the male reproductive

organ and the gynoecium represents the female reproductive organ.
MALE REPRODUCTIVE PART (ANDROECIUM)
Stamen is the male reproductive part and consists of the following parts:
a. A long and slender stalk called the filament which may be joined or free.
b. A terminal bilobed structure called the anther.
The proximal end of the filament is attached to the thalamus or the petal of the flower.
The number and length of stamens are variable in flowers of different species.
Structure of Anther
A typical angiosperm anther is bilobed with each lobe having two theca, i.e., they are
dithecous
Often a longitudinal groove runs lengthwise separating the theca.
Transverse section of an anther
It is composed of two anther sacs or lobes. Anther sacs are separated by connective
tissue.
The anther is bilobed. Each lobe consist of two theca separated by septum.
The anther is a four-sided (tetragonal) structure consisting of four microsporangia located
at the corners, two in each lobe.
The microsporangia develop further and become pollen sacs. Pollen sacs are packed with
pollen grains.

a. A typical stamen b.3D cut section of Anther

ANUSREE M NAIR
 3
II PU/CLASS XII

Structure of microsporangium:
A typical microsporangium appears circular and is generally surrounded by four wall
layers:
EPIDERMIS : It is the outermost single layer of cells which is protective in nature.
ENDOTHECIUM: it is the second layer made up of single layer of radially elongated
cells with fibrous thickening on their walls. Few cells of ensothecium in the two vertical
grooves are thin walled and constiyute Stomium, this helps in anther dehiscence.
MIDDLE LAYERS: it is the third layer composed of 1-3 layers of cells.
Endothecium and middle layers are protective in nature and help in dehiscence of anther
to release pollen grains.
TAPETUM: It is the forth and innermost layer. Cells of tapetum have dense cytoplasm
and many nuclei. It provides nourishment to the developing pollen grains.

a. T.S. of a young anther b. Enlarged view of one microsporangium showing wall layers

T.S. of mature dehisced anther with pollen grains

When the anther is young, a group of compactly arranged homogenous cells called the
sporogenous tissue occupies the centre of each microsporangium.

ANUSREE M NAIR
 4
II PU/CLASS XII

MICROSPOROGENESIS
The process of formation of microspore from a microspore or pollen mother cell by
meiosis is called microsporogenesis. Each cell of the sporogenous tissue in a
microsporangium acts as a potential pollen mother cell (PMC) or microspore mother
cell. PMC undergoes meiotic divisions to form cluster of four cells called microspore
tetrad.
On maturity anther dehydrates and the microspores dissociate from each other to develop
into pollen grains.
Inside each microsporangium several thousands of microspores or pollen grains are
formed and released with the dehiscence of anther.

POLLEN GRAIN:
• The pollen grains represent the male gametophytes.
• They develop from pollen mother cells by meiotic division.
• Pollen grains are generally spherical in shape.
• It has a two-layered wall: - hard outer layer called the exine and inner intine.
• Exine is made up of sporopollenin which is one of the most resistant organic
material. It can withstand high temperatures and strong acids and alkali. No
enzyme that degrades sporopollenin. Pollen grains are well-preserved as fossils
because of the presence of sporopollenin.
• Pollen grain exine has apertures called germ pores where sporopollenin is absent.
• The exine exhibits a different patterns and designs.
• The inner wall of the pollen grain is called the intine. It is a thin and continuous
layer made up of cellulose and pectin.
• The cytoplasm of pollen grain is surrounded by a plasma membrane.
• When the pollen grain is mature it contains two cells, the vegetative cell and
generative cell.
• The vegetative cell is bigger, has abundant food reserve and a large irregularly
shaped nucleus.
• The generative cell is small and floats in the cytoplasm of the vegetative cell. It is
spindle shaped with dense cytoplasm and a nucleus.
• In over 60 per cent of angiosperms, pollen grains are shed at this 2-celled stage.
In the remaining species, the generative cell divides mitotically to give rise to the
two male gametes before pollen grains are shed (3-celled stage).

IMPORTANCE OF POLLEN GRAINS

• Pollen grains can be used in fossil studies
• Pollen grains are nutrient rich and taken as food supplements by athletes and race
horses.

ANUSREE M NAIR
 5
II PU/CLASS XII

• Viability of pollen grains varies from species to species. Viability of polln depnd
on the temperature and humidity. In rice, wheat etc, pollen grains are viable only
for 30 minues after release. In members of Rosaceae, Solanaceae, Leguminoseae,
they remain viable for months.
• Pollen grains can be stored for many in liquid nitrogen at -196oC, to be used in
crop breeding programmes.

Pollen grains of many species cause severe allergies and bronchial afflictions in some
people often leading to chronic respiratory disorders– asthma, bronchitis, etc.
Parthenium or carrot grass that came into India as a contaminant with imported wheat has
become ubiquitous in occurrence and causes pollen allergy.

FEMALE REPRODUCTIVE PART (GYNOECIUM)

The pistil or gynoecium represents the female reproductive part of a flower. It has one or
more carpels.
When gynoecium has a single pistil, it is called monocarpellary. When Gynoecium has
more than one pistil, it is called multicarpellary:- bicarpellary (Tomato, datura) ,
tricarpellary (cucumber), pentacarpellary (Hibiscus)
The gynoecium with free carpels is called Apocarpous, E.g- Michelia The gynoecium
with carpels fused together is called Syncarpous. E.g- Papaver
• Each pistil has three parts: - stigma, style and ovary.
Stigma- It receives pollen grains. It has a sticky surface.
Style- It is the elongated slender part beneath the stigma. It connects stigma and ovary.
Ovary – It is the basal bulged part of the pistil. Inside the ovary there is a compartment
called ovarian cavity (locule).
The placenta is located inside the ovarian cavity. Ovules or megasporangium arises from
placenta.

ANUSREE M NAIR
 6
II PU/CLASS XII

The number of ovules in an ovary may be one (wheat, paddy, and mango) to many
(papaya, water melon, orchids).

THE MEGASPORANGIUM (OVULE)

• The ovule is a small structure attached to the placenta by means of a stalk called
funicle.
• Placenta is the tissue found inside ovary.
• The body of the ovule fuses with funicle in the region called hilum. Thus, hilum
represents the junction between ovule and funicle.
• Sometimes funicle extends beyond the hilum to form a ridge called Raphae.
• Each ovule has one or two protective envelopes called integuments:- inner and
outer integuments.
• Integuments encircle the ovule except at the tip, resulting in a small opening called
the micropyle.
• Basal part of an ovule opposite to micropylar end is called chalaza.
• Mass of cells enclosed within the integuments is called the nucellus. Cells of the
nucellus have abundant reserve food materials.
• Inside the nucellus is the embryo sac or female gametophyte.
An ovule generally has a single embryo sac formed from a megaspore through reduction
division.

ANUSREE M NAIR
 7
II PU/CLASS XII

MEGASPOROGENESIS:
The process of formation of haploid megaspores from the diploid megaspore mother cell
(MMC) is called megasporogenesis.
In micropylar region, the nucellus contains dense cytoplasm and prominent nucleus
which is differentiated into a single megaspore mother cell (MMC). Megaspore mother
cell is diploid and undergoes meiosis. MMC first divide transversely into two cells called
dyad. The two cells again divide transversely as a result, a linear row of four haploid cells
is produced which is called megaspore tetrad or linear tetrad.
Out of theses four megaspores, only one remains functional while the other three
degenerate.
The one functional megaspore develops into the female gametophyte or embryo sac.
This is called Monosporic development.

• The nucleus of the functional megaspore divides mitotically to form two nuclei
which move to the opposite poles, forming the 2-nucleate embryo sac.
• The two daughter nuclei undergo another mitotic division giving rise to the 4-
nucleate stage. 2 nuclei move towards the micropylar end, other two towards
chalazal end.
• The third mitotic division, which gives 8-nucleate embryo sac comprises of a
micropylar and a chalazal end with four nuclei at each end.
• Mitotic divisions are strictly free nuclear, that is, nuclear divisions are not followed
immediately by cell wall formation.

ANUSREE M NAIR
 8
II PU/CLASS XII

Organization of Embryo sac (polygonum type)

• Six of the eight nuclei are surrounded by cell walls and organised into cells; the
remaining two nuclei proceeds towards the centre of the embryo sac and are called
polar nuclei.
• Three cells are grouped together at the micropylar end and constitute the egg
apparatus.
• The egg apparatus consists of two synergids and one egg cell. The synergids have
special cellular thickenings at the micropylar tip called filiform apparatus, which
play an important role in guiding the pollen tubes into the synergid.
• Three cells at the chalazal end are called the antipodals.
• The large central cell has two polar nuclei.
Thus, a typical angiosperm embryo sac, at maturity is 8-nucleate and 7- celled.

b- 2,4 and 8 nucleate stages of embryo sac c- A mature embryo sac

POLLINATION
The transfer of pollen grains from anther to stigma of a pistil is called pollination.
Kinds of pollination:
1. Autogamy: The transfer of pollen grains from anther to stigma of same flower.
(Self pollination).
For auto gamy to occur, pollen grain release and stigma receptivity should synchronize.
Anther and stigma should lie close to each other. In normal open bisexual flowers
complete autogamy is rare.

ANUSREE M NAIR
 9
II PU/CLASS XII

2 types of flowers:-
a. Chasmogamous flowers: These are open flowers with exposed stamens and stigma
which facilitate cross pollination.
b. Cleistogamous flowers: These are bisexual closed flowers which never open at all
and anthers dehisce inside these closed flowers.
Oxalia, commelina, viola produces both types of flowers.
2. Geitonogamy: Transfer of pollen grains from the anther of one flower to stigma of
another flower of same plant. Pollens remain genetically similar.
Example: Cucurbita
3. Xenogamy: Transfer of pollen grains from anther of one flower to stigma of
another flower of a different plant. Pollens are genetically different.

AGENTS OF POLLINATION
Pollen grains cannot move by themselves requires the help of external agents to achieve
pollination. Plant need to produce more number of pollen grains.
Two main agents for pollination are:
1. Abiotic agents (Wind , Water)
2. Biotic agents (Insects, Mammals, Birds, Reptiles etc.)
Pollination by wind: It is called anemophily.
Wind pollination requires:
1. Dry, light weight, non sticky, winged pollen grains.
2. Well exposed stamens for easy dispersal.
3. Stigma should be sticky, large and feathery.
4. Numerous flowers are packed together to form inflorescence.
e.g.- corn cob tassels.
5. Wind pollinated flowers have single ovule in each ovary.
Pollination by water is called hydrophily (limited to 30 genera of monocots)
Hydrophily is common in algae, bryophytes, pteridophytes and some angiosperms.
Bryophytes and pteridophytes have less distribution as they are water pollinated.
Vallisneria and Hydrilla are submerged hydrophytes in which hydrophily observed.
Eicchornnia, Nelumbo, Water lilly etc are insect pollinated aquatic plants.
In Vallisneria (dioecious) the female flower reach the surface of water by a long stalk
and the male pollen grains are released on to surface of water. They are carried passively
by water currents, some reach stigma.

ANUSREE M NAIR
 10
II PU/CLASS XII

Zostera is a marine angiosperm where hydrophily observed: in this female flower remain
submerged in water and pollen grains are released inside the water.
In plant pollinated by water, pollen grains are long, ribbon like. It has mucilaginous
covering to prevent wetting.
Pollination by animals
Bees, butter flies, man, beetles, wasp, birds, lemur, arboreal rodents, reptiles (gecko
lizard, garden lizard) and bat are common pollinating agents.
Bees are dominant pollinating agents.

Pollination by insect is called Entomophily

Flowers of insect pollinated plants are large, sticky and brightly coloured.
They have honey and nectar glands, which are highly fragrant to attract insects.
Pollen grain surface is sticky and stigma is sticky due to mucilaginous secretion.
Polen grains get coated on to the body surface of pollinators.
Insect gets nectar and in some cases flowers provide safe place for egg laying. Example:
Amorphophallus , Yuca, Fig
Yuca and moth: Moth and plant cant complete its life cycle without each other. Moth
deposits egg in locule of ovary. Plant is pollinated and larvae comes out as seeds develop.
Some flowers produce bad odour which attract insects. Eg.- Rafflesia
Floral rewards are the benefits the pollinators gets from the plant. they are Nectar,
pollen grains and egg laying position.
Certain insects consume nectar and pollen grains, but without bringing out pollination,
they are called pollen robbers

POLLEN – PISTIL INTERACTION

All the events from pollen deposition on the stigma until the entry of pollen tube into
ovule are together called pollen pistil interaction.
It involves pollen recognition by stigma for compatible pollen.
Incompatible pollens are rejected by the pistil and compatible pollens are allowed to
grow.
The ability of the pistil to recgnise the pollen is followed by acceptance and rejection is
result of continous dialogue between pollen grain and pistil. This dialogue is mediated by
chemical components of the poolen interacting with pistil.
Pollen tube grows through stigma and style to reach the ovary. As pollen tube grows it
carriers 2 male gametes and tube nucleus. Tube nucleus guides the pollen tube.

ANUSREE M NAIR
 11
II PU/CLASS XII

It then enters the ovule through micropyle (porogamy), or through chalazal region
(chalazogamy) or there integuments (mesogamy).
Pollen tube reaches synergids, guided by filiform apparatus.

ARTIFICIAL HYBRIDIZATION:
Commonly used technique in plant breeding to obtain desirable characters.
Techniques done:
1. Emasculation: Removal of anthers from bisexual flowers before maturity of
anthers. This is done using a forceps.
2. Bagging: Emasculated flowers are covered with bag to prevent contamination of
its stigma by unwanted pollen.
3. Stigma is pollinated with desired pollens and rebagged till fruit develop.
For unisexual flowers, no need of emasculation. Flowers buds are bagged before
flowers open. Pollination is carried out with desired pollen & flower rebagged.

DOUBLE FERTILIZATION
Double fertilization is an unique feature of angiosperms.
Pollen tube grows and reach synergids, pollen tube release the two male gametes into
cytoplasm of synergid.
One of the male gamete fuses with egg nucleus to form diploid cell called zygote. This is
called syngamy.
Other male gamete fuses with two polar nuclei in the central cell to produce a triploid
primary endosperm nucleus (PEN). This is termed as triple fusion.
As syngamy and triple fusion take place simultaneously in the embryo sac, it is
termed as double fertilization.
The central cell after triple fusion form primary endosperm cell which later develops into
endosperm.
Zygote later develops into an embryo.
Double fertilization is an unique feature of angiosperms.

ANUSREE M NAIR
 12
II PU/CLASS XII

OUT BREEDING DEVICES

Most flowering plants produce hermaphrodite flowers. So pollen grains come in contact
with stigma of same flower resulting in self pollination. Self pollination result in
inbreeding depression.(leads to weaker progeny).
To discourage self pollination and increase cross pollination, flowering plants have
developed many devices this is called out breeding devices.
Contrivances or devices for cross pollination are :-
1. Dicliny (Unisexuality): When the flowers are unisexual cross pollination is rule.
2. Self - Incompatibility: In some plants, a flower cannot be fertilized by pollen of
same plant or same flower due to genetic reasons. Eg.- Solanum.
3. Dichogamy: The stamens and carpels of a bisexual flower mature at different
times.
If stamens mature earlier to carpels it is protandrous, if carpel mature early it is
protogynous.
4. In some flowers, anthers and stigmas are positioned in different positions so that
pollen cannot come in contact with the stigma of the same flower.
(Heteromorphism)

POST FERTILISATION EVENTS

The following events after double fertilization are collectively called post fertilization
events.
1. Development of endosperm from primary endosperm nucleus (PEN).
2. Development of embryo from zygote.
3. Development of seeds from ovule.
4. Development of fruit from ovary.

ANUSREE M NAIR
 13
II PU/CLASS XII

ENDOSPERM DEVELOPMENT
Endosperm develops from triploid PEN. The cells of this tissue are rich in reserve food
materials which are used for the nutrition of growing embryo.
Endosperm develops before development of embryo.
The PEN undergoes successive nuclear divisions to give rise to free nuclei. This stage of
endosperm development is called free nuclear endosperm.
Later cell wall is formed and Endosperm becomes cellular. This division is followed by
cytokinesis and endosperm is called cellular endosperm.

Functions of endosperm:
• Cells of endosperm tissue are triploid and filled with reserve food material to
nourish the developing embryo.
• Coconut water is free nuclear endosperm and the white kernel is cellular
endosperm.
• Endosperm may be completely consumed by the developing embryo. Eg:- pea,
groundnut, beans.
• In some plants endosperm may persist in the mature seeds. Eg:- Cocconut, castor.

EMBRYO
• Embryo develops at the micropylar end of the embryo sac.
• The nutrition for developing embryo is provided by endosperm.
• The zygote divides to form proembryo and subsequently to the globular, heart-
shaped and mature embryo.

EMBRYOGENY IN DICOTS
The zygote undergoes transverse division forming a large basal cell and a small apical
cell. The large basal cell enlarges and undergoes transverse division to form a group of 6-
10 cells called suspensor.
The first cell of the suspensor towards the micropylar end is called haustorium. The last
cell of the suspensor towards chalazal end is called hypophysis that later develops into
radicle.
The smaller terminal cell undergoes one vertical division. The two cells from terminal
cell divide to form four embryonal cells (quadrant stage).
Finally these four cells divide vertically forming 8-celled proembryo (octate stage).
Four cells at the apex give rise to plumule and another four give rise to hypocotyl except
its tip.

ANUSREE M NAIR
 14
II PU/CLASS XII

STRUCTURE OF DICOTYLEDONOUS EMBRYO

• A typical dicotyledonous embryo consists of an embryonal axis and two
cotyledons.
• The portion of embryonal axis above the level of cotyledons is the epicotyl, which
terminates with the plumule or stem tip.
• The cylindrical portion below the level of cotyledons is hypocotyl that terminates
at its lower end in the radical or root tip. The root tip is covered with a root cap.

A typical Dicot embryo

STRUCTURE OF MONOCOTYLEDOUS EMBRYO

• Embryos of monocotyledons possess only one cotyledon.

• Cotyledon is situated on one side (lateral) of the embryonal axis and is called
scutellum.
• The radical and the root cap are situated at the lower end of the embryonal axis in
an undifferentiated sheath called coleorrhiza.

ANUSREE M NAIR
 15
II PU/CLASS XII

• The portion of the embryonal axis above the level of attachment of scutellum is
the epicotyl. Epicotyl has a shoot apex and a few leaf primordia enclosed in a
hollow foliar structure, the coleoptile.
Eg:- Grass family, Banana, Palm etc.

SEED

Seed is a fertilized ovule. It’s the final product of sexual reproduction. It consists of the
following:
1. Seed coat- Outer integument of the ovule forms the outer seed coat (testa) and the
inner integument of the ovule forms inner seed coat (tegmen).
The micropyle remains as a small pore in the seed coat , this facilitates entry of
Oxygen and water during germination.
2. Cotyledons- 1-2 in number and rich in reserve food material. Cotyledon forms the
first set of leaves in a plant.
3. Embryonal axis.

Types of seed:
1. Non-albuminous seeds- the seed with no residual endosperm. E.g.- Pea, groundnut
2. Albuminous seeds- the seedswith residual endosperm. E.g.- Wheat, Maize, Barlry,
Castor, Sunflower.

In certain seeds nucellus may persist. This is called perisperm. E.g.- black pepper
Dormancy: In a mature seed water content decreases and seed enters a state of inactivity
called dormancy. The general metabolic activity slows down during dormancy.
When conditions ( like temperature, water, oxygen) become favourable seeds germinate.
Dehydration and dormancy of mature seeds are important for seed storage.

Advantages of seeds:
• Seeds posses better adaptive strategies for dispersal to form a new colony.
• Reserve food of seed support growth of seedling.
• They provide genetic recombination and variation as they are products of sexual
reproduction.
• Seeds can be stored for long time, this is the basis of agriculture.
• Seed coat protects growing embryo.

Seed Viability:
Vialbility is the ability of the seed to germinate. In certain species seeds lose vialbility
within few months. In some species, seeds remain viable for many years.
E.g:- Lupinus arcticus seed was excavated from Arctic tundra, estimated to be dormant
for 10000 years.

ANUSREE M NAIR
 16
II PU/CLASS XII

Phoenix dactylifera (date palm variety) – 2000 years viable seed obtained from King
Herod’s palace near Dead sea.

FRUITS
• Ovary matures into fruit and ovarian wall develops into fruit wall called pericarp.
• A typical pericarp has 3 layers: outer epicarp, middle mesocarp and inner
endocarp.
Mesocarp is fleshy in mango, mesocarp is fibrous in coconut.
• Fruits can be fleshy (orange, mango, guava) or dry (mustard, groundnut).
• Many fruits have evolved mechanism for seed dispersal.

Types of fruit:
True fruit: Fruit that develops from ovary alone. Eg:- Wheat, maize
False fruit: Fruit that develops from any other part other than ovary.
From thalamus. Eg:- Apple, Cashew.

Parthenocarpic fruit:
Fruit which are formed without fertilization are called parthenocarpic fruit.

ANUSREE M NAIR
 17
II PU/CLASS XII

These fruits do not have seeds. Eg:- In Banana parthenocarpy can be induced artificially
by growth hormones.
In Orchids, fruit contains thousands of tiny seeds. Other examples are fruits of parasitic
plants like Orobanche and Striga.
Ficus tree produces billions of seeds.
APOMIXIS
Apomixis is a form of asexual reproduction that mimics sexual reproduction by formation
of seed without fertilization.
Apomixis is observed in some species of Asteraceae and grass.
All apomictic embryos have same gentic constitution.
Methods to develop apomictic seeds:
1. A diploid egg is formed without reduction division which develops into embryo
without fertilization.
Applicaion of Apomixis : used in hybrid seed industry:-
Use of hybrid seed varieties in agriculture helps to increase the crop yield. But if the
seeds obtained from the hybrid plantys are sowm, it will not maintain hybrid characters
as segregation of characters occurs in sexual reproduction. Producing hybrid seeds each
year is costly process.
So if these hybrids are made into apomicts, there is no segregration of characters in
hybrid progeny, so farmers can keep these hybrid seeds to raise new crops each year. He
need not purchase hybrid seeds each year.

POLYEMBRYONY
The occurrence of more than one embryo in a seed is called Polyembryony. eg: Orange,
Lemon, Mango
Methods to form polyembryonic seeds:
1. Development of cells like synergids, cells of nucellus, or integument into embryo.
2. Some cells of the nucellus (diploid in nature) start dividing and develop into the
embryos and protrude into embryosac.
3. Formation of more than one embryo sac in an ovule.
4. Formation of more than one egg in an embryosac.

ANUSREE M NAIR