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BIOLOGY FIITJEE CHENNAI

CENTRE

Class Learning Improvement Program

(CLIP)Sheet with Extra notes for board
support on Chapter
Sexual Reproduction in Flowering Plants
BPVR’S NOTES

Subject: Biology

Class 12

Sexual Reproduction in Flowering Plants

It involved fusion of male and female gamete formed as a result of

gametogenesis,. The two common process of gamete formation in plants
are microsporogenesis and megaspdfogenesis found in male reproductive
(androecium) and – female reproductive (gynoecium) part of flower
respectively. The androecium is composed of stamens, while gynoecium of
carpels.
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The whole process of sexual reproduction in flowering plants can be

divided into the following four steps

1. Pollen grain formation

2. Embryo sac formation
3. Pollination
4. Fertilisation

Pollen Grain Formation

The formation of ‘ microspores or pollens is called microsporogenesis.

Pollen grain or microspore formation takes place in fertile portion of anther
called pollen sacs.
Anther is a bilobed, pollen grain containing structure. Each anther lobe is
having two chambers or pollen sacs called microsporangia, thus each
anther consists of four microsporangia.
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 A typical angiospermic anther is dithecous (i.e. having two anther lobes

and four microsporangia).
 Anther develops from a group of cells (eusporangiate development). A
young anther is made up of homogenous mass of meristematic cells
surrounded by an epidermis.
 Tapetum consists of cells with large nuclei and dense cytoplasm. The
main functions of the tapetum are the production and transport of
enzymes, hormones and food materials.
 Callase enzyme is secreted by tapetum, which dissolves callose.
 Ubisch bodies are secreted by tapetum. These are lipid in nature and
get covered with sporopollenin, which increases the thickness of exine
(i.e. outer layer of pollen wall).
 Sporopollenin present in the exine of pollen grains is resistant to
microbial and chemical decomposition.
 Pollen grain exine has prominent apertures called germ
pores, where sporopollenin is absent. ‘
 Pollen kit is an oily layer present on the outside of the mature pollen
grains of many insect pollinated species; and consists of lipids and
carotenoids.
 In microsporogenesis, the primary sporogenous cells undergo mitosis
and form diploid; miprospore mother cells or pollen mother cells or
microsporocytes. Each microspore mother cell (diploid) divides by
meiosis to form a tetrad of haploid pollen grains.
 In polyspory, more than four pollen grains are produced from one
microspore mother cell. Pollen grain is spherical Pollen kit prevents the
pollen grains by UV-radiation and 25-50 micrometer in diameter.
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Development of Male Gametophyte

 It begins inside the microsporangium. First division of cell is mitotic

producing two unequal cells, i.e. a larger vegetative cell and a small
generative cell.
 The generative cell produce two male gametes and the vegetative cell
forms pollen tube after pollination. Pollen grain at the time of pollination
may be 2-celled or 3-celled.
 In over 60% of angiosperms, pollen grains are shed at 2-celled stage
(i.e. vegetative cell and generative cell). In the remaining species, the
generative cell divides mitotically to give rise to the two male gametes
before pollen grains shed.
 Thus, in these species pollen grains shed at 3-celled stage. Pollen
grains of many species (e.g. Parthenium or carrot grass) cause severe
allergies and bronchial afflictions in some people often leading to
chronic respiratory disorders such as asthma, bronchitis, etc.
 In rice and wheat pollen grains lose viability with in 30 minutes of their
release. In some members of Rosaceae, Leguminoseae and
Solanaceae, pollen grains maintains viability for months. It is also
possible to store pollen grains of a large number of species for years in
liquid nitrogen (- 196°C).

Development of Female Gametophyte

 Gynoecium is the female reproductive part of the flower. It may be

monocarpellary (single pistil) or polycarpellary (more than one pistil).
The pistils may be fused (syncarpous) or may be free (apocarpous).
 Each pistil has three parts, i.e. the stigma, style and ovary. The
placenta is located inside the ovary. From the placent arises the
megasporangia called ovules.
 Megasporangium (ovule) is the integumented indehiscent, which
develops as a small outgrowth from the tissue of placenta. It develops
into seed after fertilisation.
 Orthotropous is most primitive and simplest type and anatropous is
most common type ovule found in angiosperm.
 The ovule is a rounded structure attached to the placenta by a stalk
called funiculus.
 The attachment place of funiculus to the body of ovule is known as
hilum.
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 Ovules are enclosed in ovary and consist of a central mass of

parenchymatous cells known as nucellus. Nucellus is bitegmic in
monocots and primitive dicots (those with crassinucellate ovules) and is
unitegmic or monotegmic as in higher dicots.
 From the base of the ovule, a third integument develops in the form of
aril in many plants namely litchi, Trianthema, Asphodelus, etc.
 A hypodermal cell from the micropylar end of ovule differentiates into a
sporogenous cell, which eventually becomes megaspore mother cell
(2n).
 The megaspore mother cell divides by meiosis to form four haploid
megaspores. One megaspore on chalazal end remains intact, while
other three degenerate.

The functional megaspore at the chalazal end grows and many vacuoles
appear in the cytoplasm.
.
Embryo Sac Formation

Out of the three nuclei, located at the micropylar end, one serves as egg
and the other two as synergids together constituting the egg apparatus.
The mature structure with two polar nuclei, antipodals and egg apparatus is
called embryo sac. In majority of flowering plants, one of the magaspores is
functional, while the other three degenerate. Only the functional megaspore
develops into the female gametophyte (embryo sac). This is called
monosporic development.

 The nucleus of functional megaspore divides mitotically to form two

nuclei, which move to the opposite poles, forming the 2-nucleate
embryo sac.
The two more sequential mitotic nuclear divisions result in the
formation of 4-nucleate and later 8-nucleate stages.
 After 8-nucleate stage, cell walls are laid down leading to the
organisation of the typical female gametophyte.
 The typical embryo sac (monosporic, 8-nucleate or Polygonum type) is
having 8 nuclei but 7 cells (3 micropylar cells, 3 chalazal cells and two
polar nuclei forming a single central cell with diploid secondary nucleus
in later stages).
 The synergids are also known as helpers. They help in distribution of
nutrients in the embryo sac with the help of filiform apparatus. They
also help in attracting pollen tube towards egg.
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 Antipodals also help in nutrition of embryo sac. A tissue called

obturator also helps in growth of pollen tube towards egg. Ovules are
located in locules of ovary.

Pollination

The transfer of pollen grains from anther of a flower to the stigma of same
or other flower is known as pollination. It is mainly of two kinds:

1. Self-Pollination
Self-pollination or autogamy or xenogamy is the transfer of pollen
grains from the anther of a flower to the stigma of the same flower or
from one flower to another on the same plant. Self-pollination produces
homozygous characters in the progeny and good characters can be as
such retained. Self-pollination is disadvantageous in the sense that no
variations are produced and the progeny becomes genetically weak.
Adaptations for Self-Pollination
Homogamy In this condition, the anthers and stigma mature at the
same time and the stigma is receptive at the time, when the pollens
shed, e.g. Mimbilis and Vinca rosea.
Cleistogamy Sometimes, the flowers do not open and remain closed
throughout their life in bud form. The pollination occurs in the bud itself.
Most of the plants with cleistogamous flowers, also bear
chasmogamous flowers,
i. e. the flowers, which open normally. Thus, cleistogamy is a facultative
character. Example of exclusively, cleistogamous flowers are very few
and include Sibularia aquatica and Polycarpon tetraphyllum.
2. Cross-Pollination
Cross-pollination or allogamy is the transfer of pollen from anther of a
flower to the stigma of the another flower, borne on separate plants of
the same species. Cross-pollination results in combination or mixing up
of characters thus, improving the quality or vigour of the species.

Agencies of Cross-Pollination

Various agencies helpful in cross-pollination are broadly categorised into

two typfts, i.e. biotic agencies and abiotic agencies.
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 Pollination with the help of air or wind is called as anemophily and

pollination taking place with the help of water is called as hydrophily.
 When biotic agents are involved in cross-pollination, the process is
collectively, called as zoophily.
 Most common type of zoophily is entomophily, i.e. pollination with the
help of various insects. Pollination through birds, is called as
omithophily, cheiropterophily is the pollination acquired through bats
and malacophily is pollination by snails.
 Myrmecophily is the name of beneficial association between ants and
flowers, in which flowers get pollinated in return of giving juicy
secretions to the ants.
 Geitonogamy is a type of pollination in which pollen grains of one
flower is transferred to stigma of another flower belonging to same
plant or genetically similar plant.

Adaptations for Cross-Pollination

Most of the flowers in nature are cross-pollinated and are thus variously
adapted for it. For avoiding self-pollination and for exclusively carrying out
cross-pollination, the adaptations are as follows:

Self-Sterility or Self-Incompatibility

In this adaptation, either the pollens of a flower are unable to grow on the
stigma of same flower or if they do so, they grow very slowly, e.g. Malva
and Abutilon.
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 Dichogamy

When maturing times of stigma and anthers is such that either stigma
becomes receptive before anthers mature (protogyny) or the anthers are
ready for dehiscence before stigma becomes receptive (protandry),
e.g. in Aristolochia and Scrophularia protogyny occurs and in rose,
sunflower, Impatiens protandry is observed.

 Heterostyly

In some members of Oxalidaceae, Rubiaceae, Polygonaceae, etc., the

flowers are dimorphic, i.e. of two different forms. One form has long
stamens and styles are very small. The anthers are well above stigma thus
minimising the chances of self-pollination. The other form has long styles
and the stamens are having small filaments. Thus, anthers lie below the
receptive part of the stigma, e.g. primrose. This condition is termed as
dimorphic heterostyly.
The cross-pollination takes place between flowers with same length of
styles. Physiology of flower also plays a role and it has been seen that
cross-pollinating flowers axe physiologically resembling to each other.

 Unisexuality (Dicliny)

The presence of only one kind of reproductive whorl in a flower is known

as unisexuality. A plant may be monoecious, i.e. carrying two different
flowers as male and female flowers on the same plant. In this case, both
cross or self-pollination can occur.
However in dioecious plants, i.e. plants in which male and female flowers
are borne on different plants, cross-pollination is the only way of pollination.

 Herkogamy

In some flowers, a mechanical barrier exists between compatible pollen

and stigma so that self-pollination is not possible. Sometimes a hood-like
covering covers the stigma as in Iris and Calotropis, the pollens are
grouped in pollinia and stick to surface till they are carried away by insects.

 Pollen-Pistil Interaction
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The pistil has the ability to recognise the suitable pollen type. If it is of the
right type, the pistil accepts the pollen and promotes post-pollination events
that leads to fertilisation. The acceptance or rejection1 of pollen by the pistil
is mediated by chemical components of the pollen interacting with those of
the pistil.
Following compatible-pollination, the pollen grain germinates on the stigma
to produce a pollen tube, through one of the germ pores. The content of
polllen grain move into the pollen tube. Pollen tube, grows through the
tissues of the stigma and style and reaches the ovary. All these events
from pollen deposition on the stigma until pollen tubes enter the ovule are
referred to as pollen-pistil interaction. In artificial hybridisation, the desired
pollen grains are used for pollination and the stigma is protected from
contamination (from unwanted pollen). This is done by emasculation and
bagging.

Fertilisation

Fusion of male and female gametes is known as fertilisation. In

angiosperms, the pollen tube has two male gametes. The phenomenon of
fertilisation was first reported by Strasburger (1884) in Monotrapa. The
pollen tube moves by its tip towards micropylar end of ovule, where egg is
situated. Male gametes are discharged in one of the synergids, which later
degenerates. Fertiliruion occurs after this and viable seeds are formed after
double fertilisation.

Double Fertilisation

It was discovered by Nawaschin in 1898 in Lilium and Fritillaria.

One male , gamete fuses with egg (syngamy) forming a diploid zygote (2n).
The second male gamete fuses with two polar nuclei (or secondary
nucleus) forming a triploid primary endosperm nucleus (3n). The process of
fusion of two male gametes in a single embryo sac is called as double
fertilisation and the formation of triploid nucleus by fusion of one male
gamete with secondary nucleus is called triple fusion. Total five nuclei
takes part in double fertilisation.
The pollen tube enters the ovule in majority of cases through micropyle by
a process termed porogamy or it may enter through chalaza (chalazogamy)
as in Casuarina and Juglans. In ‘rare cases, the pollen tube may pierce
through integuments (mesogamy) as in Pistacia and Cucurbita.
Chalazogamy was first reported in Casuarina.
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The filliform apparatus of synergids secrete some chemical substances

which direct the pollen tube towards micropyle of ovule.

Extra notes for Additional knowledge

REPRODUCTION IN FLOWERING PLANTS

INTRODUCTION
Angiosperm originated in Mesozoic era.
Angiosperm originated either in the begining of Cretaceous period or in ending
of Jurassic period of Mesozoic era. It means they are originated
between Cretaceous and Jurassic period on the earth.
Angiosperm dominated over the earth in Coenozoic era. So this era is known
as "Golden Period of Angiosperms".
First of all N. Grew realized the fact, that Stamens are male sex organ of flower
(Anatomy of plants)
Sexuality in plant first of all reported by Jacob Camerarius.
He reported Anthers are the male sex organ and Ovary with style and
stigma are female sex organ and for the formation of seed, interaction is
essential in between both the sex organs.
Significance of pollination and role of insects in pollination was recognized
by Josheph Kolreuter.
C.F. Wolf – Father of plant Embryology.
Prof. P. Maheshwari – Father of Indian plant Embryology. He wrote a book –
'An Introduction to Embryology of Angiosperms'.

Classification
Class - Dicotyledonae
Subclass - Polypetalae
Series - Thalamiflorae
Order - Parietals
Family - Cruciferae or Brassicaceae
Genus - Capsella
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Species - Bursa pastoris [Common name "Shepherd's purse"]

Most of the important Angiospermic characters are found in Capsella so that for
the study of Angiosperms, it is considered as a "Typical Angiosperm".
It is an annual plant and grows as weed during the winter season in the field.
The main plant of the Capsella is a sporophyte. Which is diploid and it is
differentiated into root, stem and leaves.
Capsella is a heterosporous plant it means there are two different types of spores
are formed in the life cycle which is classified into two categories in which male
spores are called Microspores and female spores are called Megaspores.
The process of reproduction takes place in this plant through a special structure,
called flower.
Calyx, Corolla, Androecium and Gynoecium are present in
the typical or complete flower.
The calyx and corolla are termed accessory whorls of the flower. Because these
structures do not participate in the process of reproduction, only helps.
The androecium and gynoecium are known as essential whorls, because they
are directly related with the reproduction.

Monocarpic Plants :
The plants in which flowering and fruiting takes place only once in the whole
life cycle are called monocarpic e.g. Annual & Biennial plants.

Polycarpic Plants :
The plants in which flowering and fruiting takes place many times in the entire
life cycle are known as polycarpic e.g. Perennial plant.
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Bamboo, Palms, Banana, Centuray plant (Agave Americana)

are perennial plants but they are the example of monocarpic plants.

Reproduction in Flowering Plants

"Reproduction is one of the important processes by which every living
organism make a copy of itself. It is the means of multiplication and
perpetuation of species because the older individual of each species undergo
senescence and die"
All the reproductive methods of plants are broadly categorized into two types -
Sexual Reproduction
Asexual Reproduction

Sexual Reproduction
In Angiosperms male and female gametes are formed in male and female sex
organs by the process of meiosis. Both the gametes fuse together to form
a diploid zygote which gives rise embryo. It means the process in which
embryo is formed by meiosis and fertilization is called Amphimixis.
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Male reproductive organ - Androecium

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Male reproductive organ is called androecium and their unit is called stamen.
Stamen is also known as microsporophyll. There are 6 stamens are present
in Capsella.
A typical stamen is differentiates into three parts –a long, thin structure is
called filament which joins the stamen to the thalamus. The free end of the
filament, a swollen spore bearing structure is called anther.
Anther and filament are attached together with help of small region,
called connective. Connective contains vascular tissues. The main parts of the
stamen is the anther.
Each anther generally bilobed structure i.e. anther has two anther lobes .
Each lobe of anther has two chambers (Dithecous) which are called pollen
sacs or microsporangia or pollen chambers.
Therefore, a typical anther has four pollen sacs is called tetrasporangiate.
Pollen grains are formed inside the pollen sac through the meiotic
division of pollen mother cells.
Pollen grains are formed inside the pollen sac through the meiotic
division of pollen mother cells.
At the maturity of the pollen grains, sterile tissue degenerate which are present in
between the pollen sacs. Both the pollen sac fused together. Because of this
reason, only one chamber appears in each anther lobe at maturity. So two
chambers are seen in the mature anther at the time of dehiscence.
In Capsella, which is member of the cruciferae or Brassicaceae, anther are
dithecous and tetrasporangiate type.
But in Malvaceae, the anther of stamen has only one theca in each anther lobe.
This is called monothecous and it contains only two pollen
sacs called Bisporangiate.
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STRUCTURE OF ANTHER :

The development of anther in origin is Eusporangiate type i.e. It is developed

from more then one archesporial cells.
In the transverse section of anther, it is seen almost spherical.
The following structures are present in the anther :-
Epidermis :- It is the outermost layer of anther. It is single celled thick and
continuous layer but not archesporial in origin. It forms the outermost protective
layer.
In Arceuthobium (Smallest Parastic Angiosperm) fibrous thickening present in
epidermis so it is called exothecium.
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Endothecium :- This layer is present below the epidermis. It is single celled

thick layer. During the maturation of anther, various changes takes place in
different walls of cells of endothecium. The outer wall of these cells remains thin
walled, but inner walls and radial walls become thick due to thickening of -
Cellulose fibers. Callose bands are also present along the radial walls. At some
places callose bands and fibrous thickening are absent. These places are
called stomium. The dehiscence of anther takes palce only from these
places. Endothecium becomes hygroscopic nature due to presence of fibrous
thickening. So it helps in dehiscence of anther.

Middle layer :- Middle layer is consist of parenchymatous cells. This layer

is one to three celled thick structure. Food is stored by parenchymatous cells
of this layer. Middle layer is ephemeral in nature and absent in a mature anther.
In Holoptelia plant 3 to 4 called thick middle layer is present.
In Najadaceae & Lemnaceae families middle layer is absent.
In wolffia middle layer is absent.
Tapetum :- It is the inner most layer which acts as nutritive layer. It is known
as tapetum. Pollen sacs surrounded by tapetum. This is also single celled thick
layer. The cells of the tapetum initially diploid but they become polyploidy due
to endomitosis (Chromosome duplication without nuclear division). It means
these cells contain many chromosomes and sometime binucleate
(As karyokinesis not followed by cytokinesis)
Tapetum absorbs food from the middle layer and provide nutrition to the
microspore mother cells or microspores. The cells of tapetum
secrete hormones and enzymes. The tapetum layer disappears in the mature
anther.
Note : In Nicodia and Costum plants, tapetum is multilayered.
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Pollen sacs : Four Pollen sacs are present in the anther. Pollen sacs are also
known as microsporangia. Inside the pollen sacs, microspores are formed by
the meiotic divison of microspore mother cells.
TAPETUM IS OF TWO TYPES :-
Amoeboid Tapetum/Invasive Tapetum/Periplasmodial Tapetum :
It is found in primitive Angiosperm. Such type of tapetum absorb all
foods from the middle layer. So middle layer immediately degenerates. In the
beginning, all food materials stored by tapetum. Tapetal cells convert the
absorbed food into special food granules called protoplast bodies. The
innermost layer of tapetum dissolve and release its protoplast into the cavity of
the microsporangium. Now inside the pollen sacs protoplast bodies are known
as periplasmodium. Microspore mother cells are surrounded by periplasmodium
and provides nourishment to the developing microspores. This type of tapetum
provide nutrition to the microspores after degeneration.
Example : Typha, Alisma and Tradescantia.
Glandular or Secretory Tapetum :
It is developed type of tapetum. It is not degenerates quickly. It absorbs nutrients
from the middle layer and secreted into the cavity of the microsporangia (Pollen
sacs) e.g. Usually it is found in most of Flowering plants (Capsella).
Before degeneration of cells of tapetum, they form special granules
called Proubisch bodies in cytoplasm. Proubisch bodies transfer between cell
wall and cell membrane of tapetal cells. Here they are surrounded
by Sporopollenin. Now they are called Ubisch bodies or orbicules. At last
tapetum degenerates and ubisch bodies released into pollen sacs.
Generally, sporopollenin participates in the formation of outer covering (Exine)
of Pollen grains.
Tapetum helps in transfer of food, storage of food, formation of
sporopollenin and pollenkitt materials.

MICROSPOROGENESIS :
The anther appears as outgrowth like structure in the initial stage which shows
spherical or oval shaped structure.
At this stage, it is a mass of undifferentiated and homologous meristematic
cells which is surrounded by a single cell thick outer layer. This layer is known
as epidermis. First of all vascular tissue are formed in middle region
Simultaneously four cells located just below the epidermis in vertical rows in the
region of hypodermis at the four corners are become large has visible nucleus
with dense cytoplasm. Due to this reason they are different from the rest of the
cells. These cells are called archesporial cells.
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These cells divide periclinally to form primary parietal cells below the
epidermis and primary sporogenous cells towards the centre. Both of the cells
usually undergo further divisions to form complete structure of anther except
epidermis.

Primary parietal cells undergo further periclinal and anticlinal division to form
a series of 3-5 layers making the walls of the anther.
Out of them outer most layer of anther is formed just below the epidermis by
primary parietal cells is called Endothecium or fibrous layer. The endothecium
is followed by 1-3 celled thick layer is termed middle layer. The innermost
layer of the anther which surrounds pollen sacs, is called tapetum. Later the
tapetal cells play a significant role during the meiotic cell division in
microsporogenous cells and in pollen development.
The primary sporogenous cells divide twice or more than two times by
mitotic division to form sporogenous cells and later sporogenous cells
differentiated into microspore mother cells (MMC / PMC) during the formation
of wall of pollen sac.
Each microspore mother cell divide to form four haploid microspore or pollen
grain by meiotic division or reduction division.
During this period spherical bodies are formed inside the tapetal cells before their
disintegration. These spherical bodies are known as Ubisch-body. Ubisch body is
made up of a complex substance called sporopollenin. It is biopolymer
(Heteropolysaccharide)..
After the formation of ubisch body, the tapetum layer degenerates. Ubisch bodies
participate in the formation of exine of the microspores inside the pollen sacs.
Now thick walled microspores are called pollen grains.
At the initial state all four microspores are attached together with the help
of callose layer. This group of microspores is called tetrad. After some time, this
callose layer dissolve by callase enzyme. Which is secreted by tapetum.
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Normally each microspore mother cell can form tetrad by meiotic division. But in
some plant like Zostera, some microspore mother cells become sterile and
provide nutrition to rest of microspore mother cells.
Similarly, tapetum is not well developed in Gentianaceae family so some cells
of sporogenous tissue become sterile and provide nutrition to remaining
sporogenous cells.
Types of tetrads : The arrangement of the microspores in tetrad condition as
follows :-
Tetrahedral tetrad : Four haploid microspores arranged
in tetrahedral form.
Example : Dicotyledons – (Capsella)

Isobilateral tetrad : This condition is found in monocotyledons. Microspores

are arranged at the lateral side of each other.
Decussate tetrad : In this two microspores lies at the right angle of other
microspores
Example : Magnolia
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T-Shaped tetrad : Two microspores lie longitudinally and two microspores lie at
transversly in this type of tetrad.
Example : Aristolochia & Butomopsis.

Linear tetrad : In this tetrad all four pollens arranged in linear order.
e.g., Halophylla, Halophia.
All the above type of tetrads are found in Aristolochia elegans.
Most common type of tetrads is Tetrahedral.

Structure of Microspore or Pollen Grain :

Pollen grain is the first cell of a male gametophyte.
Pollen grain is termed as immature male gametophyte. Usually, they are in
round shape. Pollen grain surrounded by two distinct layers. The outer layer
(wall) is thick, rigid and ornamented, called exine. This layer is formed
by cutin and sporopollenin. Sporopollenin is Highly resistant
material (Resistant to temperature, ph, Enzyme, Electric shock etc.) It
is nonbiodegradable.
The internal layer is thin, soft and elastic in nature. It is called intine. It is made
up of pectin and Cellulose or pecto-cellulose.

The number of germ pore, structure and ornamentation of exine is a

significant feature of taxonomy.
A detail study of pollen grains is called Palynology.
Three colpus type (slit type) of germpore are present in pollen grain of Capsella.
This type of pollen grains are called tricolpate. Only one germ pore is present
in monocots and pollen grain is called monocolpate.
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The plants in which pollination takes place by insects, their pollen grains
having oily layer around the pollen grain. It is called pollen kitt. It is composed
of lipids and carotenoids.

Function of pollen kitt :

This oily layer protects the pollen grain from the harmful ultraviolet rays.
Its sticky surface helps to attach with the insects.
Its yellow colour attracts the insects. Pollen kitt is present on the pollens
of Capsella.
Type of Germpore :

Dehiscence of Anther :
During the maturation of anther, various changes are takes place in walls of
anther.
In the begining, middle layer degenerates due to absorption of food by
tapetum.
When the micropores are formed inside the pollen sacs, at the same time ubisch
bodies are formed in cells of tapetum, then after it degenerates. Ubisch bodies
participate in the formation of exine of pollens.

In this way, in a mature anther only two layers

epidermis and endothecium are present in the form of outer covering.
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The sterile tissues are present between both the pollen sacs of each anther lobe
degenerate. So both pollen sacs of the each anther lobe fuse together to form
single pollen sac.
Therefore, in the T.S. of mature anther only two pollen sacs are
present.
Dehiscence of anther takes place during the dry season. Due to the hygroscopic
nature of endothecium loss of water takes place from the cell of
endothecium.
Walls of endothecial cells try to contract due to the loss of water
but inner and radial walls do not contract due to presence of fibrous
thickening whereas outer thin walled cells of endothecium contract and
become concave or incurved.
Incurving of outer walls creates pulling force or tension over the entire surface
of anther. Due to tension, thin walled stomial cells breaks off and dehiscence of
anther takes place and pollen grains are present in pollen sacs released into the
atmosphere.
Dehiscence of anther in Angiosperms either longitudinal or Apical pore
or Transverse or Valvular type. Dehiscence of anther
of Capsella is longitudinal.

Micro-Gametogenesis or Development of Male Gametophyte :

In flowering plants, pollen grain or microspore considered as first cell of male
gametophyte. Germination or development of pollen grain takes place before
dehiscence of anther is called precocious development. Development of pollen
also takes place at mother place [inside pollen sac of anther] is called In-situ
development.
Pre Pollination development -
In the beginning of the process, only nucleus of pollen grain divided by unequal
mitotic division, resulting two unequal size of nucleus are formed. Small
nucleus present near the walls is called generative nucleus and large
nucleus present inside the cytoplasm is called tube or vegetative nucleus.
 24

Both the nucleus surrounded by cytoplasm and it becomes dense, then again
followed by unequal cytokinesis, resulting two unequal size of the cells are
formed.
Larger cell in which large nucleus is present known as Vegetative
cell and smaller cell in which small nucleus is present, called generative cell.
Now pollen grains come in bicelled and binucleated stage. In 60%
Angiosperms pollination of pollen grains take place in bicelled and binucleated
stage. While in 40% in 3 celled, 3 nucleate stage. The development of
gametophyte take place inside the pollen grain is also known as endosporic
development.
This stage of pollen grain is called immature or partially developed male
gametophyte.
Generative cell detached from the wall and changed into vermiform or spindle
shaped structure and enter inside the vegetative cell.

Post Pollination development -

Rest of the further development of pollen grain [Immature male gametophyte]
takes place on the stigma of carpel after pollination.
Pollens absorb moisture and sugar content from the stigma. Due to this
absorption volume of internal contents of cytoplasm increased. It exerts pressure
on the both layers. Because of this pressure intine comes out through any one
germopore in the form of tube like structure called pollen tube.
 25

Facts about types of Pollen grains :

In Asclepiadaceae (Calotropis) and Orchidaceae family, all the pollen grains
joined together to form "Pollinium".
More than four pollen grains are found in tetrad called "Polyspory" e.g. Cuscuta.
In some plants, four pollens of tetrad join together permanently are called
"Compound pollens" e.g. Drosera, Typha, Drimys and Elodea.

Pollinium of Calotropis is called "Translator apparatus".

 26

Compound pollens of Mimosaceae (Mimosa) family contains 6-8 or 64 pollen

grains stick together to form a small unit that is called "Massullae"
e.g. Mimosa and Neottia plants.
Due to the presence of sporopollenin, fossils of pollen grain are always found
in good condition. The presence of fossils of pollen grains can forecast the
presence of natural resources like petroleum, coals etc. in the earth.
Pollen grains of some plants present in air cause allergy are called "aero
allergens" e.g. chenopodium, Parthenium, Sorghum and Amaranthus. ["Hay
fever" is caused by pollens of Ambrosia].
In Cyperaceae family only one pollen grain is formed from pollen mother cell
e.g., Cyperus.
Largest pollen – Mirabilis.
Smallest pollen – Myosotis.
Longest pollen – Zostera (Filiform pollen)
Eight nucleated embryo sac type of pollen is found
in Hyacinthus orientalis the pollen nucleus divide to become 8 nucleate [This
type of pollen grain discovered by Nemec. So it is called Nemec phenomenon]
Pollen grains are rich in nutrients. It has become a fashion in recent years to
use pollen tablets as food supplements in western countries. Large number of
pollen products in form of tablets and syrups are available in market. Pollen
consumption has been claimed to increase the performance of athletes and race
horse.
Pollen grains of many plant species cause severe allergies and bronchial disorders
in some people leading to chronic respiratory disorders like - asthma,
bronchitis. For ex. Parthenium (carrot / congress grass) – in India cause pollen
allergy and asthma.
Viability of pollen grains is depands on temperature and humidity. In some
cereals like rice and wheat – pollen grains loss viability within 30 minutes of
their release. In members of Rosaceae, Leguminoseae and Solanaceae pollen
grains are viable for months.
Pollen grains can be stored like semen of humen being for years
by cryopreservation in liquid nitrogen at –196º temperature.
Such pollens can be used as pollen banks similar to seed banks in crop breeding
programmes.

Female Reproductive Organ - Gynoecium

Gynoecium is the female reproductive organ. The free unit of gynoecium is
called pistil or carpel.
Carpel is also known as megasporophyll.
The carpel is differentiate into three distinct region -
 27

(i) Stigma
(ii) Style
(iii) Ovary

The free end of the carpel which receives pollen grains is called stigma. A long,
narrow tubular structure is present in between the stigma and ovary called style.
The basal swollen part of the carpel is called ovary. The ovule is also known
as megasporangia which are borne on a cushion-like tissue called placenta in
the ovary. One or more than one ovules are present inside the ovary.
The gynoecium of the Capsella is bicarpellary, syncarpous, unilocular and
superior. It becomes bilocular due to the formation of false septum or replum at
maturity.
 28

Structure of Ovule or Megasporangium :

A ridge or stalk like out growth is formed from the placenta of the ovary on
which body of ovules are present. Each ovule attached to the placenta by means
of a thin stalk called funicle or funiculus/Funiculum
Number of ovule in a ovary may be one (in wheat, Rice, Mango) or many (in
papaya, water melon orchids).
The point of attachment of the funicle with the ovule is called hilum.
The main region of the ovule is composed of mass of parenchymatous cells
called nucellus. Nucellus is the main part of ovule. The nucellus is covered
by one or two coats called integuments.
In most of the ovule, funicle is attached to the main body of ovule for some
distance (at lateral side) to form a ridge like structure known as Raphe.
Vascular tissues are present inside the funiculus which supply food
material from the placenta to the body of ovule.

Special Integuments :
ARIL – It is the type of third integuments which develops from funicle at
the base of the ovule e.g. Myristica, Asphodelus and Litchi.
ARILLODE – It develops from the tips of the outer integument and grows
downwards and surrounds the entire ovule e.g. Pitchecolobium (Inga dulce)
 29

SARCOTESTA – When outer integument becomes fleshy then it is called

sarcotesta e.g. Mangoliaceae (Magnolia).
OPERCULUM – It is a stopper or break like structure which is formed on the
micropyle. It is formed due to the elongation of inner
integuments or endostome/exostome/Nucellus projections
e.g. Lemnaceae family (Lemna).
CARUNCLE OR STROPHIOLE – It is formed due to the proliferation (out
growth) of outer integuments over the micropyle. e.g. Ricinus
communis (Castor). It is formed by sugary contents so helps in absorption of
water during germination of seeds and dispersal of seeds by ants
called myrmecochory.
COMA – In some of the plants unicellular filaments like structures are present
on the seed which is formed by cells of outer surface of outer integument. Such
seeds are known as "Comose seeds". e.g. Calotropis and Gossypium.

A place from where funicle and integuments arise is called Chalaza.

Integument is absent just opposite to the chalaza, so that a narrow passage (pore)
is formed which is called micropyle.
Micropyle in bitegmic ovule has two parts – outer region which is surrounded
by outer integument is called exostome.
The inner part of micropyle which is surrounded by inner
integument called endostome.
In most of the Angiosperm entire part of the nucellus is utilized by developing
embryo sac but in some of the Angiosperm some part of the nucellus remain
inside the ovules that part of the nucellus present inside the seed in the form of a
thin layer known as perisperm. Perisperm is commonly found in Piperaceae
(Piper nigrum) and Zingiberaceae Families (Turmeric, Ginger) and Beet plant.
Some filaments are attached with funicle [some times placenta] are known
as "Obturators".
The function of obturators is to guide the passage of pollen tube towards the
micropyle inside the ovary.

Types of Ovule on The Basis of Integuments :

A single integumented ovule is called unitegmic ovule – example – members
of Gamopetalae and Gymnosperm.
Two integumented ovule is called bitegmic ovule. Example – In most
of Angiosperm [Polypetalae – Capsella and Monocots].
The ovule in which integuments are absent is called Ategmic ovule e.g. Olax,
Liriosma, Loranthus & Santalum.
 30

Types of Ovules on The Basis of Nucellus :

Tenuinucellate – The nucellus is either less developed or present in the form of
single layer.
Example : Gamopetalae group.

Crassinucellate – The nucellus is massive type i.e., it is made up of many

layers.
Example : Polypetalae group and Monocots

The nucellus degenerates in plants of Compositae family

and integuments becomes active to form a nucellus like tissue. This is
called endothelium or integumentary tapetum. It
is multinucleate. structure.
The nucellus dissolves in the members of Podostemaceae family to form
a nutritional cavity. This is termed pseudoembryo-sac.

TYPES OF OVULES :
There are six different types of ovules are found in Angiosperms on the basis of
relationship of the micropyle, chalaza, and hilum with body of the ovule
and orientation on the funiculus :

Atropous or Orthotropous :
The body of ovule is upright in position. The micropyle, chalaza and hilum lie
in one straight line, so that this ovule is called straight or upright ovule. Example
: Betel, Piper, Polygonus and in Gymnosperms. It is the most primitive and
most simple type of ovule of Angiosperms. Raphe is absent.

Hemitropous or Hemi-Anatropous ovule :

In this ovule, the body of the ovule bent on funcile at 90º angle, i.e., body of
ovule present at right angle to the funiculus. This is intermediate
type between ortho and anatropous ovules. This ovule is also called horizontal
ovule because body of ovule present in horizontal position on the funiculus.
Micropyle and chalaza are present in the same line but micropyle is situated away
from hilum. Example: Ranunculus, Primula, Golphimia.

Campylotropous Ovule :
In this ovule, the body of ovule curved (Curvature is not effective) in this way so
micropyle and chalaza do not present in straight line. The embryo sac and
nucellus both are present in curved position. Micropyle comes close to the hilum.
Example – Leguminosae, Capparidaceae, Cruciferae family [Capsella]
 31

Anatropous Ovule :
In this type, the body of the ovule completely turned at 180º angle, due to
unilateral growth of funiculus, so it is also called inverted ovule. The chalaza and
micropyle lie in straight line. The hilum and micropyle lie side by side very close
to each other. This type of ovule is found in 80% families of Angiosperms but
not in Capsella. In this ovule micropyle facing downward condition. This is
the most common type of ovule so that it is considered as a "typical ovule" of
Angiosperms. eg. Members of Malvaceae, Cucurbitaceae, Solanaceae,
Compositae family. It is also called resupinate ovule.

Amphitropous Ovule :
In this type of ovule, curvature is more pronounced or effective in the nucellus
and due to this effect of nucellus, embryo sac becomes horse shoe
shaped. Micropyle comes close to the hilum. It is also called as transverse ovule.
e.g. Mirabilis, Lemna and Poppy, Alisma, Butomaceae family.

Circinotropous Ovule :
This type of ovule, first of all body of ovule inverted once and again turned into
straight position due to the growth of funiculus so that body of ovule present on
funicle at 360º. The entire body of ovule is surrounded by funiculus. It is also
known as coiled ovule. Micropyle is situated away from hilum e.g. Cactaceae
family-Opuntia.

MEGASPOROGENESIS :
During the development of ovule, in the begining of this
process, nucellus develops form the placenta in the form of a small rounded out
 32

growth like structure. At this stage, all the cells of nucellus

are undifferentiated and homologous and meristematic. This mass of cells
surrounded by single celled thick layer of epidermis.

Any one hypodermal cell of nucellus is differentiated and increase in size. It

becomes different from rest of the cells due to presence of distinct nucleus. It is
called archesporial cell. Archesporium divides periclinally to form an outer
primary parietal cell and inner Primary Sporogenous cell. Activity of primary
Parietal cell depends on type of plants. If plant belongs to gamopetalae then it
forms tenuinucellate type ovule and if plant belongs to polypetalae then it
forms crassinucellate type of ovule is formed. The primary sporogenous cell
directly act as a megaspore mother cell (MMC). It divides meiotically to form,
four haploid megaspores.
The four haploid megaspores generally arranged in linear tetrad. Generally the
lower most or chalazal megaspore remains functional out of tetrad of
megaspores and the other three lie towards the micropyle degenerate. This
functional megaspore produces female gametophyte. In most of Angiospersm
(Capsella), Chalazal megaspore remains functional.

DEVELOPMENT OF EMBRYO SAC OR FEMALE GAMETOPHYTE :

 33

Megagametogenesis : Megaspore is the first cell of the female gametophyte.

This megaspore grows in size and obtains nutrition from the nucellus. The
nucleus of megaspore divides mitotically to form a two nuclei. Each nucleus
moves towards the opposite pole and reached at their respective poles. Both the
nuclei lie at poles divide twice mitotically. Resulting, four-four nuclei are formed
at each poles [Total 8-nuclei].
Out of the four, one-one nucleus migrates from the both poles [one nucleus from
chalazal side and one nucleus from micropylar side] towards the centre. They are
known as polar nuclei. Both polar nuclei are present in the centre.
Remaining three-three muclei at each pole surrounded by the cytoplasm to form
cells as a result of cytokinesis. Three cells are formed towards the micropyle in
which one cell is large and more distinct out of three cells. This is called egg
cell and remaining two smaller cells are known as synergids. These three
micropylar cells collectively known as egg-apparatus. [1 Egg cell + 2
Synergids]
The three cells are formed toward the Chalaza are called antipodal cells. Both
the polar nuclei present in the large central cell. But just before the process of
fertilization they unite or fuse together in the centre to form secondary
nucleus. It is diploid in nature [2n] and one in number.
Therefore, seven cells and eight nucleated structure is formed. This eight
nucleated and seven celled structure is called female gametophyte or embryo
sac of Angiosperms. This type of embryo sac is known as polygonum
type because it is discovered by Strassburger in Polygonum plant. Polygonum
type embryo sac is most common type in Angiosperms [Capsella]. Polygonum
 34

type of embryo sac develops from single megaspore so it is also known

as monosporic embryo sac.
Fingers like processes are produced from the outer wall of the synergids are
known as filiform apparatus. With the help of these structures, synergids
absorb food from the nucellus and transfer to the embryo sac. Filiform
apparatus is less developed in antipodal cells. Filiform also secrete
some chemicals which attracts the pollen tube.
In some plants, barrier are present either above or below the female
gametophyte. These barrier are made up of thick walled cells of nucellus. They
prevent the movement of embryo sac towards the chalaza or micropyle.
The barrier which is present towards the chalaza is
called hypostase e.g. Umbelliferae
family, Zostera and Crozophora plants.
The barrier which is present toward the micropyle is
called epistase e.g. Costalia and Costum.

TYPE OF EMBRYO SACS :

Monosporic Embryo sac – It is of two types –
Polygonum type – It is eight nucleated and seven celled embryo sac.
Oenothera type – Exceptionally it is four nucleated in which only one nucleus
in a central cell and three nucleus in egg apparatus. Antipodal cells are
absent. [Micropylar megaspore become functional]
Bisporic Embryo sac – It is formed by two megaspores. It means it develop
from two nucleus of megaspores. It is of two types –
Allium type – Eight nucleated and seven celled [Chalazal megaspores]
Endymion type – Eight nucleated and seven celled [Micropylar megaspores]
Tetrasporic Embryosac – It is formed by all four megaspore
nuclei because meiosis is not accompanied by cytokinesis, so that four nuclei of
megaspores are formed.
All four nuclei are collectively known as "Coenomegaspore".
 35

POLLINATION
 36

"Pollination is defined as the process of transfer of pollen grains from anther to

the stigma of the same flower or of different flower of the same species."

Pollination is of two types :

SELF POLLINATION OR AUTOGAMY :
If the pollen grain are transferred from an anther to the stigma of the same
flower is called self pollination or autogamy.

CROSS POLLINATION OR ALLOGAMY :

When the pollen grains are transferred to the stigma of other flower of the same
species is called Cross pollination or Allogamy. It takes place in between two
different flowers.
 37

Heterostyly There is difference in between the length of the filaments of

stamens and length of style in different flowers of some plants. Some of the
plants having long stamen and short style, and in some of the plants bears long
style and short filament. Due to this reason, self pollination can not possible in
these plants e.g. Primrose (Primula), Lathyrum, Oxalis.
Primrose (Primula vulgaries) is a commonly cited example of this phenomenon.
It has two types of flowers (distyly) :
Pin-eyed or long-styled, with long style, long stigmatic papillae, short stamens
and small pollen grains.

Thurum-eyed or short styled, with short style, small stigmatic papillae, long
stamens and large pollen grains.
The stigma in the thrum-eyed flowers is at the level of anthers in the pin-eyed
flowers and vice-versa. As a rule, pollen from thrum-eyed flowers can bring
about effective pollination only in pin-eyed flowers and, similarly, pollen from
pin-eyed flowers can effect legitimate pollination in thrum-eyed flowers, but not
in their own type.
 38

Some flowers, such as Lythrum and Oxalis, show tristyly. They have three
types of flowers with respect to the length of the style and stamens. Pollen from
flowers of one type can effectively pollinate stigma of only the other two types of
flowers and not of its own type.

Self sterility or self incompatibility or intraspecific incompatibility : In this

condition the pollen grains of the flower can not germinates on the stigma of the
same flower. This condition is called self sterility. This is a parental
[Genetical] characteristic feature which is controlled by genes. Such as
in Pitunia, Malva, Thea, Passiflora, Vitis, Apple (Malus).
Incompatibility involves many complex mechanisms associated with interactions
of pollen and stigmatic tissues. It leads to prevention of pollen germination,
retardation of pollen tube growth, deorientation of pollen tube or
 39

even failure of nuclear fusion. It is controlled by genes with multiple alleles (s-
allele).

Self incompatibility or intraspecific incompatibility :

Plants rejects the self pollens grains i.e. pollens from same flower or different
flower of same plant to prevent genetic self pollination.
Recognition of "self" pollen in based on genes for self incompatibility called 'S'-
gene. In the gene pool of a plant population, there can be dozons of alleles of an
S-gene.
If a pollen grain has an allele that matches an allele of the stigma on which it
land, the pollen tube fails to grow.
Depending on species, self recognition blocks pollen tube growth by one of the
two molecular mechanism.
Gametophytic selfincompatibility (GSI)
Sporophytic self incompatibily (SSI)

Gametophytic self incompatibility – In GSI, the S-allele in the pollen genome

regulates the blocking of fertilization.
For ex. An S1 pollen grain from an S1S2 parental sporophyte will fail to fertilize
egg of an S1S2 Flower but will fertilize an S2S3 flower.
An S2 pollen grain would not fertilize either flower.
Self recognition of this type involves the enzymetic destruction of RNA within a
rudimentary pollen tube RNA hydrolyzing enzymes are secreted by style of
carpel which destroy pollen tube.
Ex. Liliaceae, Poaceae, Solanaceae, Trifolium.

Sporophytic self in compatibility (SSI) – In this system pollens of a plant

behave similarly, irrespective of S-allele they carry.
For ex. A plant carrying S1S2 alleles, the pollen carrying S1 or S2 allele, would
behave as S1 if S1 is dominant or as S2 if S2 is dominant and S1 + S2 if no
dominance. So the presence of even one of the alleles of the stylar tissue in
sporophytic tissue of male parent make the all pollen of that plant non-functional
with respect to that particular style.
So this S1S2 plant produce pollens S1 or S2 are completely incompatible to plant
carrying S1S2, S1S4, S1S5 or S2S3, S2S4, S2S5 but 100 percent compatible with a
plant carrying S3S4 or S3S5
In SSI system rejection occurs on the stigma the pollen tube is destroyed as can
not penetrate cuticle of stigma.
Ex. Astaraceae, Brassicaceae
 40

Growth of its pollen tube of self pollinated pollen grain is very slow and growth
of the pollen tube of cross pollinated pollen grain in very fast so pollen tube of
cross pollinated pollen grains reach earlier inside the ovule. This is
termed prepotency.