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Class - Dicotyledonae
Subclass - Polypetalae
Series - Thalamiflorae
Order - Parietales
Family - Cruciferae or Brassicaceae
Genus - Capsella
Species - bursa pastoris [Common name "Shepherd's purse"]
Angiosperm orginated in Mesozoic era.
Angiosperm originated either in the begining of Cretacious period or in ending of Jurassic period of Mesozoic
era. It means they are originated between Cretacious and Jurassic period on the earth.
Angiosperm dominated over the earth in Coenozoic era. So this era is known as "Golden Period of
First of all N.Grew realized the fact, 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'.





Locule Accessory
sex organs


Ovule Thalamus

Placenta Pedicel


E 1
Most of the important Angiospermic characters are found in Capsella so that for the study of Angiosperms, it
is considerd 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
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 accessary 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
According by Goethe, Flower is modified shoot which shows favourable adaptation for reproduction
through a special method. Flower has a small or long stalk like structure called pedicel. Free end of the
pedicel is flattaned or dome shaped is called thalamus. The thalamus is a type of modified stem, on which
nodes and internodes are present.
Nodes are present very close to each other and internodes are small highly reduced in the thalamus.
The whorls present in the flower are the modifications of leaves and arranged in four circles on the thalamus.
The four nodes are present on the thalamus, in which first modified leaves are attached on the first lower
node are called calyx.
The corolla born on the second node. Androecium are present on the third node and gynoecium on the
fourth node in uppermost position.
In some of the plants the length of internode increases which is present in between Calyx and Corolla is called
Anthophore e.g. Silene plant, Dianthus.
The length of internode between the corolla and androecium increases. It is called androphore e.g. Passiflora.
If the internode between androecium and gynoecium increases then it is called gynophore. e.g. Capparis.
Both androphore and gynophore are present in the same flower is called gynandrophore or androgynophore.
e.g. Gynandropsis pentaphylla and Cleome gynandra.
Calyx is also modified vegetative leaves. Such as in Mussaenda flower, one sepals of calyx modified into
leaf like bright and attractive yellow colour structure called "Advertising flag". It helps in pollination
In some flowers; thalamus grows inside the ovary is called carpophore e.g. Coriandrum and Foeniculum.
In Nymphaea Petals like stamens are present.
On the basis of above examples, we can prove "Flower is modified shoot".
The plants in which flowering and fruiting takes place only once in the whole life cycle are called monocarpic

e.g. Annual & Biennial 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.
Bamboo, Palms, Banana, Centuary plant (Agave americana) are perennial plants but they are the example
of monocarpic plants.

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"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 categorised into two types -
1. Sexual Reproduction 2. Asexual 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. The process in
which embryo is formed by meiosis and fertilization is called Amphimixis.

Male Reproductive Organ Androecium

Male reproductive organ is called androecium and their unit is called stamen.
Stamen is also known as microsporophyll. There are 6 stamens 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 Connective
are attached together with help of small region, called connective. Connective
contains vascular tissues. The main parts of the stamen is the anther . Filament

Each anther generally bilobed structure i.e., anther has two anther lobes A B
is called dithecous.
Each lobe of anther has two chambers which are called pollen sacs or pollen chambers.
Therefore, a typical anther has four pollen sacs is called
Connective Callose bands
Pollen grains are formed inside the pollen sac through Microspore
Middle layer
mother cells
the meiotic division in pollen mother cells.
Stomium Microsporangia
At the maturity of the pollen grains, sterile tissue
Anther wall
degenerate which are present in between the pollen Endothecium
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 T.S. of anther, showing four
mature anther at the time of dehiscence. microsporangia and anther walls
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 anther lobe. This is called monothecous and it
contains only two pollen sacs called Bisporangiate.
Monothecous anther are also found in Moringa, Wolffia plants.
In Arceuthobium where there is only one microsporangium per anther. This condition is called


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 :-
(i) 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.

E 3
In Arceuthobium (Smallest Parasitic Angiosperm) fibrous thickening present in epidermis so it is called
(ii) 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.
Note: In Hydrocharitaceae family fibrous thickening is absent in endothecium.
(iii) Middle layer :- Middle layer consists of Endothecium
parenchymatous cells.This layer is one to three celled Epidermis Middle layers
thick structure. Food is stored by parenchymatous
cells in this layer. Middle layer is ephemeral in nature
and absent in a mature anther.
Note : (1) In Holoptelia plant 3 to 4 celled thick
middle layer is present. (2) In Najadaceae &
Lemnaceae families middle layer is absent. (3) In
Wolffia middle layer is absent.
(iv) Tapetum :- It is the inner most layer which acts as Microspore
nutritive layer. Pollen sacs are surrounded by tapetum. mother cells
This is also single celled thick layer. The cells of the Stomium

tapetum initially diploid but they become polyploid T. S. of young anther showing four
and multinucleate due to endomitosis, free nuclear microsporangia and vascular bundle.
division and polyteny. It means these cells contain
many chromosomes.
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.
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 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.

It is advanced type of tapetum. It does not degenerates quickly. It absorbs nutrients from the middle layer and
secretes into the cavity of the microsporangia (Pollen sacs) and does not store it 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.
4 E
Note - Tapetum helps in transfer of food, storage of food, formation of sporopollenin and pollenkitt
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 division of microspore mother cells.
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 homogenous 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 group of 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.
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 Primary
Archesporial cell Epidermis parietal cells sporogenous cells

Meristematic Procambial
cells strand
T. S. of young anther Differentiated Differentiated primary parietal
archesporial cells cells and primary
microsporogenous cells

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 by mitotic division to form sporogenous
cells and later sporogenous differentiated into microspore mother cells 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. Ubischbody is made up of a complex substance called sporopollenin. It
is the polymer of carotenoids.

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 stage 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
Note :- 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.
E 5
Note :- Similarly, tapetum is not well developed in Gentianaceae
family so some cells of sporogenous tissue become sterile and Microspore mother cell
provide nutrition to remaining sporogenous cells.
Types of tetrads :- The arrangment of the microspores in tetrad
Successive Simultaneous
condition as follows :-
type type
(i) Tetrahedral tetrad :- Four haploid microspores arranged In Monocot Microsporogenesis In Dicot

in tetrahedral form
Example :- Dicotyledons- (Capsella).
(ii) Isobilateral tetrad :- This condition is found in
monocotyledons. Microspores are arranged at the lateral
side of each other.
(iii) Decussate Tetrad :- In this two microspores lies at the
right angle of other microspores
Example :- Magnolia
(iv) T-Shaped tetrad :- Two microspores lie longitudinally
and two microspores lie at transversly in this type of tetrad.
Example :- Aristolochia & Butomopsis.
(v) Linear tetrad - In this tetrad all four pollens arranged in A B

linear order. e.g., Halophylla, Halophia.

All the above type of tetrads are found in Aristolochia
elegans. MOTHER CELL.
Most common type of tetrads is Tetrahedral.

Tetrahedral Isobilateral Decussate T-Shaped Linear


Facts about types of Pollen grains

(1) In Asclepiadaceae (Calotropis) and Orchidaceae family, all the pollen grains joined together to
form "Pollinium". Pollinium of Calotropis is also called "Translator apparatus".
(2) More than four pollen grains are found in tetrad called "Polyspory" e.g. Cuscuta.
(3) In some plants, four pollens of tetrad join together permanently are called "Compound pollens" e.g.
Drosera, Typha, Drimys and Elodea.
(4) 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.
(5) 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
(6) Pollen grains of many plants are rich in nutrients. It has becomes a fashion in recent years to use pollen
tabletes as food supplements. In western countries, a large number of pollen products in the form of
tablets and syrups are available in the market. Pollen consumption has been claimed to increase the
performance of athletes and race horses.

6 E
(7) In Cyperaceae family only one pollen
grain is formed from pollen mother Translator
locating paired Gynostegium
cell. e.g., Cyperus.
pollinia Corpusculum
(8) Largest pollen - Mirabilis. (Adhesive, bilobed)
(9) Smallest pollen - Myosotis. Translator locating
Paired pollinia
(10) Longest pollen Zostera (Filiform
pollen) Disc

(11) Eight nucleated embryo sac type Pollinia

of pollen is found in United
Caudicle style
Hyacinthus.[This type of pollen (Retinaculi 2)
grain discovered by Nemec. So it is (HYgroscopic)
called Nemec phenomenon] A C B
(12) Dumble shaped pollens are found in


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 resistent material. It
is nonbiodegradable.

Due to the presence of sporopollenin, fossils of pollen grain

are always found in good condition. The presence of fossils of STRUCTUREOF POLLENGRAINS
pollengrains can forecast the presence of natural resources
like petroleum, coals etc. in the earth.

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.

Usually, at few places on outer surface exine is absent or present in the form of thin layer. These thin places are
called germ pore. The intine comes out through the any one germpore during the germination of pollen grain
in the form of pollen tube.

The number of germpore, 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 most of the dicots (Capsella). This
type of pollen grains are called tricolpate. Only one germ pore is present in monocots and pollen grain is
called monocolpate.

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 :-
(i) This oily layer protects the pollen grain from the harmful ultraviolet rays.
(ii) Its sticky surface helps to attach with the insects.
(iii) Its yellow colour attracts the insects. Pollen kit is present on the pollens of Capsella.

E 7
Type of Germpore :-


During the maturation of anther, various changes are takes place in walls of anther.

In the beginning, 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, bundle Epidermis
then after it degenerates. Ubisch bodies participate in the Pollen sac

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.
Stomium Pollen
The sterile tissues are present between both the pollen sacs grains
of each anther lobe degenerate. So both pollen sacs of the
each anther lobe fuse together to form single pollen sac. Dehiscence of Anther

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 exert 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.

Longitudinal Porous Transverse Valvular

Capsella Solanum Ocimum sanctum Berberis

8 E
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. So it is called as precocious
development. Development of pollen also takes place at mother place [inside pollensac of anther] is called
In-situ development.
(i) Pre Pollination development
In the beginning of the 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 wall is called generative
nucleus and large nucleus present inside the cytoplasm is called Tube or Vegetative nucleus.
Both the nucleus surrounded by cytoplasm and it becomes dense, then 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 Angiosperms pollination of pollen
grains take place in bicelled and binucleated stage in 60% Angiosperms and in 40% Angiosperms
pollination occurs at 3-celled stage. In this generative cell divides and form two male gametes. The
development gametophyte take place inside the sporangia 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.
(ii) Post Pollination development
Further development of pollen grain [Immature male gemetophyte] takes place on the stigma of
Carpel after pollination. Pollens absorb moisture and sugar content from the stigma. Due to this volume
of internal contents of cytoplasm increased. It exerts pressure on the both outer layers. Because of this
pressure intine comes out through any one germpore in the form of tube like structure called pollen
First of all vegetative nucleus enter into the pollen tube and assumes terminal [tips] position. This
spindle shaped generative now enter into the pollen tube. Inside the pollen tube, generative cell divides
mitotically and to form a two non motile male gametes. Now male gametophyte comes in three
celled structure in which one vegetative cell and two male gametes are present.
This three celled stage represents the mature male gametophyte of Angiosperm [Capsella
also]. Male gametophyte is highly reduced and completely depends on sporophyte.
First of all pollen tube discovered by G.B. Amici in Portulaca plant.
Longest pollen tube is found in Zea mays.

Development of male gametophyte with the fomation of male gametes.

E 9
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 -
[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 ovules is also known as megasporongia which are borne on a
cushion-like tissue called placenta in the ovary. One or more than Ovule
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. A typical

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
The point of attachment of the funicle with the ovule is called hilum.
The main region of the ovule is composed by 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 Chalazal end
3 Antipodals
like structure known as Raphe.
Vascular tissues are present inside the Nucellus
2 Polar nuclei
funiculus which supply food material from
the placenta to the body of ovule.
A place from where funicle and
Egg cell
integuments arise is called Chalaza. (Female gamete)
Micropylar end
Integument is absent just opposite to the
chalaza, so that a narrow passage (pore) (A) Mature ovule and (B) Mature embryosac
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 nuclellus 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)

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.
(i) ARIL It is the type of third integuments which develops from funicle at the base of the ovule
e.g. Myristica, Asphodelus and Litchi.
(ii) ARILLODE It develops from the tips of the outer integument and grows downwards and surrounds
the entire ovule e.g. Pithecolobium (Inga dulce)

10 E
(iii) SARCOTESTA When outer integument becomes fleshy then it is called sarcotesta e.g. Magnoliaceae
(iv) 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. e.g. Lemnaceae family (Lemna).
(v) 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.
(vi) 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 single integumented ovule is called unitegmic ovule - example - members of Gamopetalae and
Two integumented ovule is called bitegmic ovule. Example - In most of Angiosperm [Polypetalae-Capsella and
The ovule in which integuments are absent is called Ategmic ovule e.g. Olax, Liriosma, Loranthus &
Santalum .
(i) Tenuinucellate - The nucellus is either less developed or present in the form of single layer.
Example :- Gamopetalae group.
(ii) 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 polyploid structure.
The nucellus dissolves in the members of Podostemaceae family to form a nutritional cavity. This is
termed pseudoembryo-sac.
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:
[i] ATROPOUS OR Micropyle

The body of ovule is upright in Chalaza

position. The micropyle,
chalaza and hilum lie in one C
straight line, so that this ovule A Micropyle B Amphitropous
is called straight or upright Orthotropous Anatropous

ovule. Example :- Betel,

Piper, Polygonum and in
Gymnosperms. It is the most
primitive and most simple
type of ovule of E F
Angiosperms. Raphe is Campylotropous Hemitropous Circinotropous
absent. Different type of Ovules

E 11
In this type, the body of the ovule completely turned at 1800 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.
In this ovule, the body of the ovule bent on funcile at 900 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.
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 nucellus is present in curved position but the embryo sac remain
straight. Micropyle comes close to the hilum. Eg.:- Leguminosae, Capparidaceae, Cruciferae family
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 , Poppy, Alisma, Butomaceae family.
In this type of ovule, body of ovule inverted and again turned into straight position due to the growth of
funiculus so that body of ovule present on funicle at 3600. 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.
During the development of
ovule, in the beginning of
Primary Primary
this process, nucellus parietal cell sporogenous cell
develops form the placenta Primary Parietal cell
archesporial cell
in the form of a small
rounded out growth like mother cell
structure. At this stage, all
the cells of nucellus are
homogenous and

meristematic. This mass Linear tetrad of

megaspores Functional
of cells surrounded by
single celled thick layer
of epidermis. Different type of Megasporogenesis

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
12 E
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. The primary sporogenous cell
directly act as a megaspore mother cell. 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 Angiosperms [Capsella], Chalazal
megaspore remains functional.


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
Remaining three-three nuclei at each pole surrounded by 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
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 nautre [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 embryosac of Angiosperms. This type of embryosac is known as
"polygonum type" because it is discovered by Strasburger in Polygonum plant. Polygonum type embryosac
is most common type in Angiosperms [Capsella]. Polygonum type of embryosac develops from single
megaspore so it is also known as monosporic embryosac.
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 embryosac. Filiform
apparatus is less developed in antipodal cells. Filiform also secrete 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 embryosac towards the chalaza or micropyle.
The barrier which is present towards the chalaza is called "hypostase" e.g. Umbelliferae family and
Zostera and Crozophora plants.
The barrier which is present toward the micropyle is called "epistase" e.g. Costalia and Costus.
(i) Monosporic Embryosac It is of two types

1. Polygonum type It is eight nucleated and seven celled embryosac.

2. 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. Endosperm will become diploid. [Micropylar
megaspore become functional]
(ii) Bisporic Embryosac It is formed by two megaspores. It means it develop from two nucleated
megaspores. It is of two types

E 13
1. Allium type Eight nucleated and seven celled [Chalazal megaspores]
2. Endymion type Eight nucleated and seven celled [Micropylar megaspores]
(iii) 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 colletively known as "Coenomegaspore".
Generally all the four Nuclei of megaspores take part in the formation of the seven celled and eight
nucleated embryosac. Beside this different other types of embryosacs are also formed. Arrangement
of nuclei are not definite in embryo sac in some of Angiosperms. So that different type of embryosacs are
formed as follows :-

Types of Tetrasporic Embryosac Number of nuclei in Sec.nucleus Ploidy of endosperm

1. Adoxa type 2 3n
2. Chrysanthemum type 3 4n
3. Plumbago type 4 5n
4. Plumbagella type 4 5n
5. Fritillaria type 4 5n
6. Penaea type 4 5n
7. Peperomia type 8 9n

"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 :-
If the pollen grain are transferred from an anther to the stigma
of the same flower, or different flowers of the same plant is
called self pollination or autogamy.

When the pollen grains are transferred to the stigma of other


flower of the same species is called Cross pollination or

m y

Allogamy. It takes place in between two different flowers.

Cross pollination is of two types :-
(i) Geitonogamy :
po Se
llin lf
When, pollination takes place in between the two flowers at
of the same plant then it is called geitonogamy. From
the genetical point of view geitonogamy is self
pollination because all flowers of the same plant are


genetically identical. But ecologically, it is considered as



p oll

cross pollination.

(ii) Xenogamy :
When the pollination takes place in between the two
different flowers of two different plants of the same
species then it is called xenogamy. This is real or true B
cross pollination. Genetically, as well as ecologically,
it is cross pollination. Pollination

14 E
Contrivances or Adaptation for Self Pollination :
(i) Monocliny (Bisexuality) - It means flowers are bisexual.
(ii) Homogamy :- When both the sex organs of a flower
mature at the same time. It is called homogamy. It Normal
increases chances for self pollination. E.g. Mirabilis,
Catharanthus .
(iii) Cleistogamy :- In some plants bisexual flower are formed
which never opens throughout the life. Such flowers are
called cleistogamous flowers, such as Commelina, Underground
Viola, Oxalis, Juncus, Drosera. Commelina plant
have two types of flowers. One type of flowers are
cleistogamous and another are chasmogamous
A special type of cleistogamy is found in flowers of legume
plant. The sex organs are closed in a structure which is Commelina plant
formed by joining of some petals. It is called keel. This
keel never opens . So only self pollination takes place in
these plants.
(iv) Bud pollination:- This pollination occurs in bud stage before the opening of flowers. E.g. Pisum,
Wheat, Rice.
Contrivances for Cross Pollination
(i) Dicliny (Unisexuality) :- Presence of unisexual flowers confirm cross-pollination. Self pollination never
takes place in these flowers. It means allogamy becomes compulsory.Examples Morus, Palms, Date
Palms, Cucumber and Cucurbita, Carica.
(ii) Dichogamy :- In many bisexual flowers of the plants, stamens and carpels of a flower do not mature
at the same time. Dichogamy is of two types
(a) Protandry :- The anther of a flower mature earlier than carpels, is called protandry. Many plants of
Angiosperms are cross pollinated only because of protandrus condition. E.g. Salvia, Sunflower,
(b) Protogyny :- The carpels of the flower mature earlier than stamens. It occurs in few plants e.g. Ficus
bengalensis, Saraca indica, Ficus religiosa, Aristolochia, most of plants of Cruciferae and
Rosaceae family .
(iii) Chasmogamy or Anthesis :- Opening [blooming] of the floral bud in the form of a flower is called
(iv) Herkogamy :- In some plants, morphological barriers are formed in between the anther and stigma of
the same flower, so self pollination can not occurs and pollen grains from the anther are unable to reach
the stigma of the same flower. In this condition, only cross pollination is possible. e.g. Gloriosa, Calotropis
and caryophyllaceae family.
(v) Heterostyly :- There is difference in between the length of the filaments of stamens and length of style
in 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, Linum, Primula.
(vi) 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. If the incompatibility is due to the genotype of the sporophytic/stigmatic tissues, it is termed
sporophytic incompatibility on the other hand, if it is due to the genotype of the pollen, it is termed
gametophytic incompatibility. This may be due to prevention of pollen germination, retardation
of growth, deorientation of pollen tube or even failure of nuclear fusion. It is controlled by genes
with multiple alleles (s-allele). Enlarged pollen tube turns upwards and degenerates in style.

E 15
(vii) Prepotency: Growth of its pollen tube of self pollinated pollen grain is very slow and growth of the
pollen tube of cross pollinated pollen grain is very fast so pollen tube of cross pollinated pollen grains
reach earlier inside the ovule. This is termed prepotency. e.g, Apple, Grapes



When the pollen grains are transfer from one flower

to the another flower through the air is called
anemophily and flower is known as anemophilous
flowers. Such as Cereal plants :- Maize etc.
Anemophily is also found in all Gymnosperms. The
anemophilous plants produce enormous amount of
pollen grains. The pollen grains are very small, light
weight and dry and their stigma is hairy or brushy
and mucilagenous (Sticky).
m al e
i nf
Yellow clouds are formed in the sky during the wind lor
e sc
pollination in Pinus. These yellow clouds are formed nc
Styles e
due to the pollen grains are called "Sulphur

Anemophilous flowers are neither attractive nor with

fragrance. They do not have nectar glands.
Anemophilous flowers are generally unisexual.

E.g. - Sugar cane, Bamboo, Coconut, Cannabis,

Grasses, Date palms, Typha, Oak, Hazel,
Air pollination in Maize plant
Cucumber, Papaya, Cotton, Tobacco,Eucalyptus.

Potamogeton and Myriophyllum are aquatic plant but it is anemophilous.

Maximum loss of pollen grains takes place only in this pollination. It is completely non directional process.

Female Male
When the pollination brings by water flower
is known as hydrophily. It is of two

(i) Epihydrophily :- When the

pollination takes place on the
surface of water is called
epihydrophily e.g.
Vallisneria and Ruppia.

(ii) Hypohydrophily :- When the Stigma


Anther lobe
pollination takes place inside
the water is called
hypohydrophily. e.g.
Ceratophyllum, Najas Female Male
flower flower
Zostera and Hydrilla .
Specific gravity is found in A inflorescence B C
pollen grains so they remains
Epihydrophilly in Vallisneria
suspended in water.

16 E
When the pollination brings by animals is called zoophily. Generally in zoophillous plants, flowers are very
large, attractive and maximum number of nectar glands are present (more than entomophillous).
The pollination takes place with the help of insects is known as entomophily. Most of insect pollination (80%)
only by Honey bees. Most of entomophilous plants are ornamental plants. Ornamental plants utilize their
maximum energy in this pollination and develops different types of adaptation for attraction of insects. These
flowers are attractive in colour. They posses special fragrence. Nectar glands is also present. e.g. Rose,
Lemon. The pollen grains of insect pollinated flowers become sticky due to presence of pollen kitt. The
surface of stigma of flowers is rough.
Some of the plants develop special adaptation for insect pollination.

In Mussaenda plant, advertising flag is present for attraction of insects.

The lever mechanism or turn pipe

mechanism is found in Salvia for pollination.
The flowers of Salvia have bilabiatecorollate
tube. The connective of stamen is long. The
anterior anther lobe of connective is fertile while
posteriorly lobe is sterile. When the insect lands
on the lower lip, the fertile lobe automatically
brings down to touch the back of insect and
thus depositing the pollen grains on the back of

Some of the flowers have attractive bracts i.e.

bright and coloured e.g. Bougainvillea
Yucca plant has develops an obligate C
symbiotic relationship with an insect- Pronuba
moth. The pollination in "Yucca takes place Lever mechanism in Salvia
only by Pronuba moth. This insect lays eggs in
the ovary of flower. Life cycle of both depend
on each other.



Male flower

A B Female flower Gall flower Pronuba

Aristolochia Ficus Yucca

"Trap door mechanism" is found in species of Ficus (Peepal) for pollination [Blastophaga insect] because
Hypanthodium type of inflorescence is present.
In Aristolochia "Fly trap mechanism" is found for pollination. This flower is known as "Pit fall flower".

E 17
Ophrys speculum flower pollinated by Wasp [Colpa aurea] by means of pseudo-copulation. The
appearence and odour of the flower like female wasp [Mimicry].
In Centaurea plant pollination takes place by "piston mechanism".

In Rafflesia, the pollination is brought about by Carrion flies and dispersal of seeds by Elephant.

Nymphaea, Nelumbo and Alisma are also entomophilous plants while they are hydrophytes.
(ii) Ornithophily :-
Then process of pollination is taken place by birds is known as ornithophily.
e.g. Sun bird or humming bird in Bignonia plant and by Honey bird in
Strelitzia, Callistemon (Bottle brush), Bombax [Silk cotton tree,] Butea
(iii) Chiropterophily :-
If the pollination brings through the bats (Pteropus) is called
chiropterophily. These flowers are big in size e.g. Anthocephalus
kadamba, Bauhinia, Kigelia plants, Adansonia, Musa. Humming bird

(iv) Myrmecophily :-
This pollination brings about by Termites and Ants. e.g. Prosopis (kikar), Acacia (Mimosoideae
family), Mango and some members of Rubiaceae family.
(v) Malacophily or Malmacophily :-
This pollination brings about by Snails e.g., Aspidistra lurida and Chrysanthemum plants.
Erythrina (Coral tree) plant visited by Crows and Squirrels.
In Arisaema or Cobra Plant pollination by snake called "Ophiophily".

Night flowering plants are pollinated by Moths. They are highly scented.Their flower generally white coloured

Orchids are pollinated by Wasps/Snails.

Favourable colour of Honey bees is yellow, but they are blind to red colour.

Larger animals such as some primates (lamurs), arboreal (tree-dwelling) rodents, or even reptiles (gecko lizard
and garden lizard) have also been reported as pollinators in some species.

The fusion of male gamete with female gamete is called fertilization. First of all, fertilization was
discovered by Strasburger (1884) in Monotrapa plant. This process is completed in the following steps :-
After pollination, pollen grains germinate on the stigma. They absorb moisture and sugar contents from
stigma and swellup. The intine of pollen grain grows out through the any one germinal pore of exine, in the form
of tube like out growth is called pollen tube. One pollen tube develops in Capsella and most of Angiosperms
is called monosiphonous condition, but more than one pollen tubes develops in Malvaceae and Cucurbitaceae
family. It is called polysiphonous.

When the pollen tube comes down from the stigma into the style, first of all vegetative nucleus enter, into the
pollen tube then it is followed by generative cell. The tube nucleus always occupies in terminal position in pollen
tube. The vegetative nucleus controls the growth of the pollen tube. Mean while, the generative cell divide
mitotically to form two male gametes. Both of the male gametes are non motile.
Boron and calcium ions (mainly Boron) are essential for the growth of pollen tube and best temperature for
growth of pollen tubs is 20300C. Pollen tube shows apical growth and chemotropic movement.

18 E
Finally, the pollen tube enters in the ovary at that time, ovule becomes mature. Inside the ovary obturators
guides the passage of pollen tube towards the micropyle. A mature ovule in which embryo sac also matured, has
three paths for the entry of pollen tube:-
(i) POROGAMY :- In this, pollen
tube enters into the ovule through
Pollen tube
the micropyle. It is known as
porogamy. It is found in most of
Angiosperms [Capsella].
(ii) CHALAZOGAMY :- In this Pollen tube

method, the pollen tube enter into Pollen tube

the ovule through the chalaza. This Porogamy Chalazogamy Mesogamy

method is di s c o v e r e d in
Casuarina by Treub [1891] e.g. Various methods of entry of pollen tube into the ovule
Betula and Juglans (walnut).
(iii) MESOGAMY :- In this method, pollen tube enter into the ovule either through
integuments- Cucurbita or through the funiculus - Pistacia and Populus.
Pollen tube can enter into the ovule through the any passage but inside embryosac, it enter only through the egg
apparatus. After the entrance inside the ovule, it grows towards the egg apparatus because synergids cells
secrete the chemical (hormones) which attracts the growth of pollen tube. It means pollen tube shows
chemotropic movement in ovule.
Any one synergid starts degenerating when the pollen tube comes near egg apparatus. The pollen tube enter
into the embryosac through the degenerating synergids.
When tip of the pollen tube enters into the embryosac vegetative nucleus degenerates. The tip of the pollen
tube swells and burst [Endosmosis] after reaching inside the embryosac. The pollen tube released all contents
including both male gametes inside the degenerating synergids of embryosac.
Two dark granules appears in the region of degenerating synergids. These are known as
X-bodies. They are two in no. and both X-bodies are formed by the degenerating nucleus of tube cell and
Before or after the entrance of pollen tube into the embryosac, both polar nuclei of the central cell fused together to
form a diploid nucleus. It is known as secondary nucleus or definitive nucleus.
Out of two, one male gamete fertilized with egg cell and to form a diploid zygote. This fusion is known as
syngamy. This is true mechanism of fertilization process.
The second male gamete fused with diploid secondary nucleus which is formed by the fusion of two polar
nuclei. This fusion is known as triple fusion resulting, a triploid (3n) structure is formed. It is called primary
endosperm nucleus.

Fertilization takes place twice at a time in Angiosperm is called double fertilization.

Double fertilization was discovered by "Nawaschin" in Lilium and Fritillaria plants.

Double fertilization and triple fusion is the specific or universal characteristic of Angiosperm. There
are five nuclei and three gametes participate in double fertilization.
A zygote is formed by true fertilization (syngamy] develops into embryo. Triploid primary endosperm nucleus is
formed by triple fusion develops into the endosperm which is used as nutrition for growing embryo.

E 19
All the remaining cells of embryosac like antipodal cells, synergids degenerate excluding zygote and primary
endosperm nucleus after the fertilization. At this time, zygote obtains food from degenerating synergids and
antipodal cells.

The fertilization in which non motile gametes are carried to female gamete through pollen tube is known
as "Siphonogamy".

Entry of more than one pollen tube into the ovule leading to occurence of super numerary male gametes is
called "Polyspermy."

Pollen tube bursts
and releases 2 sperm
Polar nuclei

Egg cell

Triple fusion 2 fused

polar nuclei + sperm 3x Primary
endosperm nucleus

Double fertilisation
Antipodal and
synergids degenerated
(Egg cell + Sperm) 2x Zygote

Release of male gametes and double fertilization

First of all endosperm develops from the primary endosperm nucleus after the fertilization which stored food
materials. It is utilized by the embryo during the early development then after at the time of seed germination.
Food is present in the form of starch in endosperm. The endosperm is of three types on the basis of
development :-

This type of endosperm mostly found in Dicotyledon [Polypetalae]. Nuclear endosperm is also present in
Capsella .
Such type of endosperm develops by free nuclear divisions of nucleus of primary endosperm nucleus, Thus
a multinucleated endosperm is formed. Later on cytokinesis takes place, so that multicellular endosperm is

This type of endosperm is the most common in Angiosperms.

The milky fluid is found in green Coconut is example of nuclear endosperm, which is called liquid syncytium.

In Melastoma, cytokinesis never take place so that it is always remains nuclear endosperm.


This type of endosperm is found in Gamopetalae group. During the development, each division of primary
endosperm nucleus is followed by cytokinesis. So that endosperm is remains cellular from the biginning.

20 E

During the development of this type of endosperm, first division of primary endosperm nucleus is followed by
unequal cytokinesis so that two unequal sized cells are formed (Cell towards the micropyle is large). Now free
nuclear divisions takes place in each cells, results it becomes multinucleated. Eventually cytokinesis takes place
later on so that it is changed into a cellular endosperm. This type of endosperm is found in Monocots. It is
intermediate type of endoperm.

Micropylar end embryo

Nuclear type


Cellular type

Endosperm Embryo

Helobial type

Different types of endosperms in Angiosperms

After the observation of first two divisions of primary endosperm nucleus, endosperm can be identified whether
it would be nuclear, or cellular endosperm.
Endosperm is absent in some of Angiosperms e.g. In Orchidaceae, Podostemaceae and Trapaceae.
Exceptionally, some of the plants have diploid endosperm instead of triploid such as in Oenothera.
Maize and Tomato have mosaic endosperm in which patches of different colours are present.
The endosperm in Betalnut is rough surfaced. It is known as "ruminate endosperm".
The drinking portion is nuclear endosperm and edible portion is cellular endosperm in Coconut.

Development of embryo in Capsella is first time discovered by "Hanstein".


In Angiosperm, Zygote undergoes in resting phase. When the endosperm is formed, development of zygote starts.
In the beginning it absorb food from the endosperm and increase in size then after a layer secreted by itself. Now
it is called Oospore.
The first division of Oospore is transverse, results two cells are formed. The one cell lies towards micropyle
is called basal cell or suspensor cell. The other cell is formed towards the Chalaza is called apical cell or
terminal cell or embryonal cell.
The basal cell and embryonal cell divide simultaneously.

E 21
The basal cell divides transversly and apical cell divides vertically resulting, two suspensor cells and two embryonal
cells formed. This stage is made up of four cells which are arranged in 'T' shaped structure. Embryonal cells
divide vertically to form four Embryonal cells. This is the quadrant stage of embryo.

The two suspensor cells divided by transverse divisions forming a 6-10 celled long filament like structure is
termed suspensor. The main function of suspensor is to pushes the developing embryo into food laden endosperm
to provide nutrition.

The micropyler cell of the suspensor swells up. This cell of suspensor is known as haustorial-cell.

The cell of suspensor lies near the embryonal cells is called hypophysis. This cell combined with radicle to
form the apex of root [Root cap].

These four cells quadrent embryo further divide transversely to produce eight. The eight celled stage of embryo
is called octant stage. The eight cells of octant are arranged in two tiers.

The four cells of the octant embryo lies near the hypophysis is known as hypobasal cells and four cells
present towards the chalaza termed epibasal cells.

The hypobasal cells gives rise to radicle and hypocotyl and epibasal cells giverise to two cotyledons and
plumule of the embryo.

All the cells of octant divided by periclinal division so that a 16 celled globular embryo is formed [Proembryo].

Due to fast division of embryonal cells of globular embryo, a heart shaped embryo is formed. All the cells of
this embryo are meristematic.

Due to the fast growth in two lobes of heart shaped embryo, they develop into two cotyledons. Both the
growing cotyledons turn in downwards due to the curved position of body of ovule of Capsella.
The tissues are present above the joining place of both the cotyledons are responsible to form plumule and
behind it epicotyl is formed.
The tissues present opposite to the plumule give rise to radicle.
This curved position of the embryo is called Torpedo or Chordate stage.
An axis is present between plumule and radicle is called embryonal axis. It is also called Tigellum [main
embryonal axis].
Both the cotyledons are present at lateral position of embryonal axis and plumule is formed in terminal position
in Dicotyledon embryo.
This type of development of embryo is known as Crucifer type or Onagrad type. It is the most common
type of development in Dicots.
Crucifer type of development is also found in Capsella, so that it is considered as typical Angiosperm for
the study of embryonic development of Angiosperms.
During the development of embryo, embryo is formed from the some part of zygote so that is called meroblastic

Ovule modified into seed in which testa is formed by outer integument and tegmen is formed by inner
Only micropyle of ovule remains unchanged and also present in seed.

Entire ovary modified into fruit. This fruit is formed by fertilized ovary so that it is called true fruit.

In some of the Angiosperms fruit is formed from the ovary without fertilization known as parthenocarpic fruit.

In some fruit parthenocarpy is useless (If edible part is endosperm or seed)

22 E

Terminal cell

Basal cell

Haustorial cell


Root cap
Suspensor Cotyledons
Haustorial cell




The Lilium type of embryonic
Cotyledon cell
development is found in Embryonal cell

monocotyledons. The first division is

Embryonal axis cell
transverse division in Oospore. Results
two cells are formed the upper cell Zygote Oospore Suspensor Vesicular suspensor
chalazal is called embryonal cell and cell
Apical cotyledon
lower micropylar cell is termed as basal Cotyledon
cell. The basal cell does not divide further Plumule initial
and later on it increases in size and form Coleoptile
Radicle initial
single celled vesicular suspensor. Plumule
Only embryonal cell divides transversly in
which terminal cell is called cotyledon Coleorhiza
cell and lower (Middle) cell is known as

embryonal axis cell.

A transverse division takes place in
Development of Embryo in Monocotyledon
embryonal axis cell to gives rise two cells.
The one cell out of two, gives rise to plumule initial and another gives rise to radicle initial.
The plumule initial divides to form a plumule of the embryo.
Radicle initial divides to form radicle. In this both the initials are responsible to form embryo in Lateral position.
An apical cotyledon is formed by the continuous divisions of cotyledon cell.

E 23
In Angiosperms, development of embryo is meroblastic and Scutellum
endoscopic [towards the chalaza].
In the grass family the cotyledon is called scutellum that is situated towards
one side (lateral) of the embryonal axis. At its lower end, the embryonal
axis has the radical and root cap enclosed in an undifferentiated sheath Epiblast
called coleorrhiza. 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 Radicle
Root cap
coleoptile. Coleorrhiza
AAL.S. of an embryo of grass

Development of floral organs in Arabidopsis

Arabidopsis thaliana is a small weed belonging to family Brassicaceae. It contains approximately 26000
genes and the complete gene sequence has been studies by plant biologists. Like other plants in Arabidopsis the
initiation of flowering is influenced by various environmental factors, Of these photoperiodism and temperature are
more important. Under the influence of the two factors the apical merisem starts producing flowers instead of vegetative
structur. The development of calyx, corolla, androecium and gynoecium is controlled by specific genes. These are
called homeotic selector genes, e.g. Apetala-2, Apetala-3, Agamous etc. As a result of effect of these genes, the
development of leaves is replaced by the development of flowers.

Similar to leaves, the floral organs developed by periclinal divisions in protodermal or subprotodermal cells.
Initially these cells show periclinal divisions followed by periclinal as well as anticlinal divisions resulting in formation of
an outgrowth. Cells of the outgrowth divide, develop and differentiate to form different floral organs. In Arabidopsis
calyx are the first to develop followed by corolla, androecium and gynoecium. The order of development of floral may
vary from plant to plant. For example in membes of family Umbelliferae the sequence of development of floral
organs is - androecium, corolla, calyx and gynoecium.



Petal Petal



Development of floral organs in Arabidopsis

(A) intial stage of development (B) fully developed flower

24 E
Difference between Self and Cross pollination
S.No. Self pollination Cross pollination
1. Pollen grains are transferred to stigma Pollen grains are transferred to stigma of
of same or genetically similar flower. genetically different flower.
2. Anthers and stigma mature simultaneously. They mature at different time i.e. protandry
and protogyny.
3. It occurs in open as well as closed flower. It occurs in only open flower.
4. It is very economical for plants. It is not economical as the plant has to produce
large number of pollen grains, nectar, scent
and colouration .
5. External agencies are not required. It is essential i.e. depends on agencies.
6. Young ones are homozygous. Young ones are heterozygous.
7. It produces pure lines because of the non It produces variations due to genetic
occurrence of genetic recombinations. recombinations.
8. It cannot eliminate harmful traits. It can eleminate harmful traits.
9. Useful characters are preserved. Not preserved.
10. No adaptability in the changing environment. It provides adaptability.
11. It cannot introduce new traits. It can introduce new traits.
12. Disease resistance is low. Disease resistance is optimum.
13. There is decrease in yield. It increase the yield.
14. It does not help in the development of new species. Helps in the development of new species.

Difference between Embryo and Endosperm

S.No. Embryo Endosperm
1. It is formed by fertilized egg (syngamy) It is formed by fusion of secondary nucleus
(triple fusion)
2. It is always diploid structure It is generally triploid structure
3. It gives rise to new plant It provides nutrition to the developing embryo.
4. Cotyledons, plumule and radicle are Such type of structures are never formed.
formed in embryo
5. Embryo is present in seed. It is only found in endospermic seeds otherwise
It degenerates with the formation of seed.
6. Embryo dies in the absence of endosperm Endosperm does not die in the absence of embryo.
7. Germination of seed takes place. There is no germination is found in endosperm.

Difference between Egg cell and Secondary nucleus

S.No. Egg cell Secondary nucleus
1. It is present near the micropyle, inside It is present in the middle of the embryosac

the embryosac.
2. Generally egg is surrounded by two synergids. It is not surrounded.
3. Only single nucleus is present in it. It is formed by fusion of two polar nuclei.
4. It is haploid structrue. It is diploid structure.
5. It is fertilized with one male gamete It is fused with one male gamete and to
and to form a diploid zygote (Embryo) form a triploid primary endosperm nucleus

E 25
Difference between male and female gametophyte
S.No. Male Gametophyte Female Gametophyte
1. It is developed from microspore or pollen grain. It is developed form megaspore.
2. It does not remain embedded permanently It remains embedded permanently
in microsporangium. in megasporangium.
3. Male gametes come out of pollen grain Female gamete always remains inside,
due to the formation of pollen tube. covered by membrane of megasporangium.
4. There are two phases of growth - Only single phase of growth.
prepollination and post pollination.
5. It is three celled structure in mature stage. It has seven cells in mature stage.
6. It will disintegrate after fertilization. Two new structures are formed after
fertilization, that is endosperm and oospore.
Difference between Epigeal and Hypogeal germination
S.No. Epigeal germination Hypogeal germination
1. Cotyledons come above surface of soil Cotyledon remains under the soil surface.
during the germination and remains attached
till the formation of new leaves.
2. Due to the growth of hypocotyl, plumule Due to very less growth of hypocotyl, cotyledons
and cotyledon pushed above the ground. do not come above the ground.
3. Growth of epicotyl does not take place. Due to high growth of epicotyl, plumule
comes above ground and cotyledon remains
inside the ground.
4. Cotyledons become green and perform Cotyledon remains under the soil and do not
leaf like function perform leaf like function
5. Cotyledon is responsible to form first Cotyledon decomposed inside the soil.
leaves, than after dries and fall on the ground


1. Zygote 2n
2. Embryo 2n
3. Endosperm 3n
4. Radicle 2n
5. Plumule 2n
6. Cotyledon 2n
7. Nucellus 2n
8. Integument 2n

9. Microspore mother cell 2n

10. Megaspore mother cell 2n
11. Ovary wall 2n
12. Carpel 2n
13. Sepal, Petal 2n
14. Stamen 2n
15. Leaf, Root, Stem 2n

26 E
S.No. Parts of Plants Number of Chromosomes
1. Microspore n
2. Tube cell n
3. Male gamete n
4. Megaspore n
5. Embryosac n
6. Synergid n
7. Antipodals n
8. Egg cell n
9. Polar nuclei n+n

Difference between pollination and fertilization

S.No. Pollination Fertilization

1. Transfer of pollen grains from anther to Fusion of male gametes and egg cell in
stigma of the flower is called pollination. embryosac situated inside the ovule is
called fertilization.
2. This process take place before fertilization. This process take place after pollination.
3. For the completion of this process, insects, There is no any external medium is utilized in
water, air like agencies are essential this process
4. Pollen tube is not formed Pollen tube is formed which transfer male
gametes up to egg cell.
5. This process take place on outer parts This process take place inside the flower,
of flower, so that it is external mechanism so that it is internal mechanism.

Difference between Monocotyledonous and Dicotyledonous seed

S. No. Monocotyledonous Seeds Dicotyledonous Seeds

1. Only single cotyledon is present with Two cotyledon are present with embryo
2. Generally cotyledon is thin or papery Cotyledons are thick
3. Generally seeds are endospermic Generally seeds are non endospermic, some times
may be endospermic
4. Cotyledon is also called scutellum Not called by this name
5. In seed plumule is covered by coleoptile Coleoptile and coleorhiza are not formed.
and radicle is covered by coleorrhiza.

6. Plumule is in lateral position and Plumule is in terminal position and cotyledons

cotyledon are is in terminal position are present in lateral position
7. Radicle degenerates after sometime and Radicle is responsible to form primary root.
adventitious roots are formed at that place.
8. In some of the seeds, seed coats and Such types of seed are not found.
cotyledon fused together e.g. Wheat etc.

E 27

Angiospermic plant


Carpel (Microsporophyll)
Seed (Megasporophyll)

Pollen sac
Ovule (microsporangium)
Sporophyte (Megasporangium)
Embryo (2n)

Megasporogenous Microsporogenous
Tissue Tissue

Zygote Microspore
Megaspore Mother cell
Mother cell

Syngamy Triplefusion Reduction division


Sec. Nucleus Megaspore

(2n) (Pollen grain)
Female Gamete
(Oosphere or egg) Female
Gametophyte Male
(Embryosac) Gemetophyte

Male Gemete


28 E



Developing embryo

Main plant body

Zygote Microsporangia

Primary endosperm
nucleus Sporophytic pahse

Gametophytic phase
(x) Meio
(redu s
Double ction is
al div
fertilisation ision
Microspore tetrad

Egg cell

Sperms in
pollen tube

Mature male 4 megaspores


3 megaspores

Mature female
Functional megaspore

E 29
In asexual reproduction, the new individual are produced by any means other than the fusion of sex gametes.
It means a "reproduction in which new individuals are formed without meiotic division and fusion of
gametes is called asexual reproduction." In this way asexual reproduction is also known as apomixis.
[Greek - Apo = without; mixis = mixing] Apomixis term suggested by Winkler.
The Apomixis is characterised by quick multiplication and reproduction of genetically similar plants from the
single parent. Such a population produced from single individual is called "clone" and each member of the
clone is called ramet.
In flowering plants, there are two main types of Apomixis.
(i) Agamospermy
(ii) Vegetative Propagation

In this type of method embryo is formed without fertilization and meiotic division is called agamospermy. It
means plants belonging in this category propagated through seeds but the embryo formation does not involve
meiosis and syngamy.
There are three different type of agamospermy -
In this method archesporium differentiates to form a megaspore mother cell but this megaspore
mother cell directly gives rise to an embryosac without meiosis. This embryo sac is diploid and a diploid
embryo is formed without fertilization from diploid egg of this embryosac.
Example : Parthenium, Taraxacum
Parthenogenesis : In this process haploid egg cell of female gametophyte is responsible to form
a haploid embryo without fertilization.
Apogamy : In this process any haploid cell of female gametophyte except egg cell is responsible
to form a haploid embryo without fertilization.
Parthenogenesis and Apogamy both are not included in agamospermy.
Note : If both gametophyte and sporophyte are diploid in Parthenogenesis and apogamy then it
is called diploid Parthenogenesis and diploid apogamy.


In this method, an embryo is formed from any diploid cells [Nucellus or integuments] of the ovule
except embryosac. This diploid cell behave like a zygote. Adventive embryony derived from Nucellus in
Citrus, Mangifera, Opuntia, Mamillaria and from Integuments in Spiranthus australis.
It is dicovered by Rosenberg in Heiarcium plant. In this method embryosac or female gametophyte is
directly formed from any diploid cell of the sporophyte except megaspore mother cell without meiosis
is known as apospory. In this gametophyte always remains diploid, e.g. Heiracium, Ranunculus
and Rubus.

Many embryos are formed inside the single seed is called polyembryony. first of all, it is observed by
Leeuwenhoek in Citrus (Orange) seeds. Polyembryony is commonly found in Gymnosperms but it is
also found in some of Angiospermic plants such as Orange, Lemon and Nicotiana etc.
If polyembryony develops naturally then it is called spontaneous polyembryony [self]. When it is
developed artificially is called induced polyembryony.
When many embryos are formed from separate-separate embryosacs [more than one] inside the
ovule is called false or pseudo polyembryony.

30 E
When many embryos are formed inside the single embryo sac of the seed is called "true poly
embryony". It is either develops from :-
(i) Clevage of Zygote or budding - e.g. Cymbidium, Exocarpus, Nymphaea, Nicotiana,
Orchids etc.,
(ii) By the fertilization of synergids e.g., Anemone, Aristolochia, Sagittaria etc.
(iii) Fertilization of antipodal cells - e.g., Paspalum, Ulmus etc.
Note : In Lilium all three types of polyembryony are present.
[Adventive embryony is also example of polyembryony in which additional number of embryos
are formed from nucellus or integuments]
This concept was given by Haberlandt and practically proved by Steward. It is based on totipotency.
Anther of the plant grown on culture medium by Guha and Maheshwari. As a result of this culture,
haploid and diploid two different type of plants are formed. Diploid plants developed from the wall of
the anther and haploid plants developed from the pollen grains. Such type of haploid plants
obtained from the tissue culture are known as androgenic plants.
Fast vegetative multiplication through the tissue culture from any part [cell or tissue] of the plant in the
field of Agroforestry and Horticulture is called "micropropagation".
(i) P.Maheshwari established Research centre of Angiosperm Embryology at Delhi
(ii) Book published by Maheshwari - An Introduction to the Embryology of Angiosperms.
Plants belonging to this category propagate by a part of their body other than a seed. The structural unit that is
employed in place of seed for the propagation of new plants is called propagule. In Angiosperms any parts
of the plants - roots, stems and leaves used for vegetative propagation.
Generally method of vegetative propogation have been further divided into two types-
(a) By roots :- Modified tuberous root of Sweet potato (Ipomoea batatas), Asparagus, Tapioca,
Yam, Dahlia and Tinospora can be propagated vegetatively when planted in soil.
In some plants adventitious buds develop on the ordinary roots like-Dalbergia sisso, Populus,
Guava, Murraya, Albizzia lebbek etc. which grow to form new plants.
The buds present on the roots grow into leafy shoot above the ground is called Slips.
(b) Under ground Stem :- In some plants under ground modified stem such as -
Rhizomes - Typha; Canna, Ginger, Turmeric, Lotus, Musa etc.
Corm - Gladiolus, Colocasia, Crocus, Amorphophallus, Alocasia etc.
Bulbs - Onion, Garlic and Lilies
Tubers - Potato, Helianthus tuberosus etc. which grow to form a new plants.
Sucker - Mint and Chrysanthemum;

(c) Creepers :
In creeping stem of the plants adventitious root are developed from the nodes and to form a aerial
shoots such as
(i) Runners - Cynodon, Oxalis and Centella
(ii) Stolon = Fragaria (Strawberry) & Vallisneria
(iii) Offset - Pistia, Eichhornia (water hyacinth) etc
Aerial stem - Opuntia

E 31
(d) Leaves :
Some plants produce adventitious buds on their leaves e.g., Bryophyllum, Begonia,
Streptocarpus, Saintpaulia. These buds remains dormant, when the leaves attached with
plants but after separation, when it comes in contact with moist soil develop new plantlets [buds]
which form new plants.
In Kalanchoe plant, whole portion of leaf blade regenerate a new plant.

In some of the plants, fleshy axillary buds develop from axis of leaves are called Bulbils
Example Dioscorea, Oxalis, Dentaria, Globba, Agave, Lilium.
There are special type of fleshy buds develop in Aquatic plant are called Turions.
e.g. Potamogeton, Utricularia.
Gardeners and Horticulturist have employed the various method of vegetative propagation for economic
production. All the methods are man made so that their practices constitute artificial means of vegetative
propagation. These are as follows-
[i] Cutting :- A cutting is separated portion of root, stem or leaf which is used for propagation, Factors are
taken into consideration for successful propagation such as (i) age of the parent plant, (ii) length and
diameter of the cutting, (iii) season and (iv) type of plants. Some time the stem cuttings are treated with
rooting hormone [IBA, IAA or NAA] for proper development of adventitious roots e.g., Sugar cane,
Rose, Croton, Tapioca, Chinarose (Stem cutting) Lemon, Tamarind (Root Cutting),
Sanseviria (Leaf cutting). [Favourable time for cutting - Rainy season]
[ii] Grafting : - It is the most common method of vegetative propogation described by ancient gardeners
long before the science of horticulture became established. In this method, parts of two plants are joined
in such a way that they grow as one plant. Grafting is done between two closely related dicotyledonous
plants having vascular cambium. The rooted supported portion of one plant called Stock is joined
with a twig of another plant called Scion. Generally, the root stock belong to wild variety which is
resistant to disease & pest. The scion is derived from the plant possessing better characters. In
grafting, the stock and scion cuts are marked in such a oblique manner so that they fix with each other.
This joint covered with clay or a layer of wax. Within a few days, tissues of stock and scion combined
together and to form a new plant. e.g., Grafted Mango, Roses, orange, Seedless Grapes and
Guava, Apple, Pear. [Favourable time for grafting - Spring season]
Types of Grafting -
(a) Tongue/Whip grafting (b) Wedge grafting
(c) Crown grafting (d) Side grafting (e) Approach grafting
Bud grafting - This method is similar to grafting except that scion in this case consist of a bud along with
some portion of bark having intact cambium.
Generally bud grafting is done during the rainy season e.g., Roses, Peach.


Scion, fitted Stock
Stock over stock

Tongue Wedge Crown Side

Various types of grafting

32 E
[iii] Layering :- This method is used in those plants which are having
flexible long branches. In this method, roots are artificially induced
on the stem branches before they are detached from the parent plants
for propagation. There are two common types of layerings.
(a) Mound/Ground Layering - This method is only utilized in
herbacious plants. In this method, the lower branch of stem is
bent down and partially defoliated and injury is made on defoliated
part. It is covered by thin layer of moist soil in such a way that the
growing tip of the branch remains above the soil surface. After a Plaster of dung,
few days the pegged portion develops adventitious roots. Then clay, hay and
rooted branch cut and separated from parent and grown into a
new plant e.g., Jasmine, strawberry, Raspberry etc.
(b) Air Layering or Gootee - This method is commonly employed in case of shrubs and trees
which do not possess branches near the ground. In this method a ring of bark is removed [girdled]
from the aerial branch. This girdled portion is covered with moist grafting clay [ 2parts clay +
1 part cow dung + some pieces of hay + cotton + water] and wrapped with a polythene sheet.
This wrapped portion of the branch is called gootee. Inside the gootee roots developed within a
period of a or two month(s) e.g., Litchi, Pomegranate, Orange, Lemon, Bougainvillea,
Guava etc.
According to P. Maheshwari :
Recurrent apomixis Non recurrent apomixis

Agamospermy Vegetative Parthenogenesis Apogamy


(a) Merits of Vegetative propagation :

(i) It is good for the multiplication of seedless plants e.g. Banana, Sugarcane, Pineapple and seedless
Orange and Grape.
(ii) It is the fastest method of reproduction e.g., Potato crop requires more than one year with the help
of seeds, however, it takes only 3 to 4 months with the help of tubers. Similarly, Lily takes 4 to 7 years
through seeds, however, 1 to 2 years by bulbs.
(iii) By grafting, desirable quality of fruit/flower/seed can be obtained.
(iv) Disease free plants can be cultured by microprogagation and micrografting.
(v) Plants with long seed dormancy or poor seed viability or poor seed can be propagated vegetatively
e.g., Cynodon dactylon (Lawn, Doob or Bermuda grass).
(vi) Good quality and better yield varieties can be preserved for a long duration in offsite collection,
herbarium, botanical gardens etc.
(vii) It gives 100% genetical similarity to their parents i.e., clone.

(b) Demerits of Vegetative propagation :

(i) Diseased parents always give diseasd clone.

(ii) Clone undergoes degeneration due to the absence of sexual reproduction.

(iii) Vegetative organs can not be preserved for long duration like culms of Sugarcane.
(iv) Vegetative propagatory organs or structures cannot be safely and easily stored as comparison to
seeds. They are easily decomposed and attached by various pathogens like Bacteria, Virus, Fungi.
(v) Do not cause any variation in plantlets, thus decrease in the adaptation power.
(vi) There is absence of dispersal mechanism. Vegetative propagation in a particular area causes
overcrowding. It results in intraspecific competition.

E 33
1. It is the only method of reproduction in those plants which have lost their capacity to produce seeds -
e.g. Banana, Seed less grapes, Orange etc.
2. The plants which produce small quantities of seeds, poor viability of seed or prolonged seed
dormancy, reproduce only by this method because it is more rapid, easier and less expensive.
3. Production of these plants more as compared to seeded plants.
4. The greatest advantage of this method is that a biotype of plant [Original plant] can be retained and
multiplied indefinitely without any change or variation.


Asexual Reproduction Sexual Reproduction

1. It does not involve the formation and fusion It involves the formation and fusion of two gametes,
of gametes. generally distinguished into male and female.
2. New individual develops from one cell or a New individual develops from zygote or fusion
part of one parent. product of two gametes, which may or
may not be produced by two parents.
3. New individuals are genetically similar to the New individuals or offsprings are generally
parents. different from either of the two parents.
4. It involves only mitotic divisions. It involves meiosis at one or the other stage.
In higher plants, it occurs at the time of
5. It does not require the formation of sex Formation of sex organs is a pre-requisite for
organs. sexual reproduction.
6. It does not introduce variability. Hence, It introduces variability and is, hence of evolutionary
has no evolutionary importance. importance.

Young plant
Young shoot
New plants
eye bud
Adventitious buds New plant
Old root tuber New root tubers


Reproductive root Aerial shoots arising from the

buds in the regions of nodes
or eyes in Potato tuber.
Sweet Potato and Dahlia

34 E


Fleshy root

Bulbil Adventitious
A leaf of Bryophyllum showing
Bulbils : A. Dioscorea ; B. Oxalis ; C. Agave sp. formation of new plants from
marginal adventitious buds.


Tongueing Stem Ringing



New plant Roots

Inter node
Adventitious roots
Stem Notching

A portion of Sugarcane A bud growing into

Stem having buds new plant


Vegetative propagation by layering

Adventitious roots

Polythene covering

Grafting clay

Ring of bark

Approach grafting
Air layering (Gootee)

E 35


Bud scion
Callus Embryoid inserted in
bark of stock
Plantlet T-shaped
Nutrient slit
solidified Bark of scion
with agar A


Micropropagation : A. Culture tube with Bud grafting

callus B. Embryoids ; C. Plantlets


The whole plant exists in a seed. Plant is the main base of whole living world. So it can be say that seed is
the most important structure. In angiosperm, after the fertilization, egg cell changed into embryo, ovary
changed into fruit and ovule change into seed.
Morphologically, ripened ovule is known as seed. In other words, seed is a mature, integumented
Seeds are characteristic of Spermatophytes (Gymnosperms and Angiosperms). Typical mature seed is having
three main parts :
1. Seed coat :
Outer, protective covering of the seed is called seed coat, which develops from integuments of ovule. In seeds
developing from bitegmic ovules, there are two distinct layers in seed coat. The outer layer is thick, hard and
leathery (developing from outer integument), called testa, whereas inner layer is thin and papery or
membranous (developing from inner integument), called tegmen. In seeds developing from unitegmic ovules
there is single layered seed coat. The seed coat performs usual protective function.
The seed is attached to the fruit wall or pericarp by means of a stalk called funicle or funiculus. The point of
attachment of funiculus to the body of mature seed is called hilum. A small opening or pore called micropyle
is present just near the hilum, which is the way of entry of water into the seed. As most of the ovules in

angiosperms are inverted or anatropous, so a ridge called raphe is present.

2. Embryo :
Embryo is the most important part of the seed, which represents mini or tiny future plant. Embryo develops
from fertilized egg [zygote].
The embryo is having an embryonal axis or main axis called tigellum, to which one or two cotyledons (seed
leaves) are attached, depending upon whether the seed is monocot or dicot. The portion of embryonal axis
or tigellum below the point of attachement of cotyledons, is called hypocotyl, which bears radicle or future

36 E
root at its tip. Similarly, portion of embryonal axis or tigellum above the point of attachment of cotyledons,
is called epicotyl, which bears plumule (future shoot) at its tip.
In some seeds (e.g. legumes), reserve food is stored in cotyledons, whereas in others (e.g. cereals), there is
special nutritive tissue called endosperm. The seeds having endosperm are called endospermic or
albuminous seed, e.g., Cereals, Castor, etc., whereas seeds in which endosperm is fully consumed by
embryo and no endoserm is left, are called non-endospermic or ex-albuminous seeds, e.g., Gram, Pea,
Sem, Cucumber, Tamarind, etc. The reserve food materials in seeds may be carbohydrate (e.g., Wheat,
Rice) or proteins (legumes) or fats (Castor, Peanut, Sunflower), etc.
Some time, some part of nucellus remains unused which is present in the form of thin layer around the
endosperm is called perisperm. e.g. Betel nut, Black pepper, Castor.
In Monocot embryo, there is a single cotyledon called scutellum. e.g. Grasses like Wheat. [On opposite
side of scutellum is a tongue shaped outgrowth remains of second cotyledon is present called epiblast.]
In Dicot albuminous seed-castor (Ricinus communis), there is a specific outgrowth called caruncle or strophiole,
present over micropyle. It is formed by proliferation of cells of outer integument at tip. Caruncle is somewhat
spongy and helps in absorption of water during germination of seed.
On the basis of cotyledon in the seed of angiosperm are called Monocotyledon [one cotyledon] and
dicotytedons [Two cotyledon].
3. Endosperm :
It is the nutritive tissue which may present or absent in the seeds. The angiospermic seed classified into two
categories on the basis of presence or absence of endosperm in seeds -
1. Non Endospermic or Ex-albuminous seed 2. Endospermic or Albuminous seed.
1. Non Endospermic or Ex-albuminous seed :
Such type of seeds do not have an endosperm at maturity, therefore are called non endospermic or Ex-
albuminous seeds. The endospermic tissues are absorbed during the development of embryo. The absorbed
food materials from the endosperm is stored in cotyledons, that why they become so large and fleshy e.g.
Capsella and most of dicotyledons. But Castor seed is endospermic.
Non-endospermic dicot seeds : e.g., Gram, Bean, Pea, cucumber ,Tamarind.
Non-endospermic monocot seeds : Pothos (money plant), Vallisneria, Alisma, Amorphophallus.

2. Endospermic or Albuminous seed :

This type of seeds, food is stored in endosperm. Such seeds are called endospermic seed OR Albuminous
seed. The endospermic tissue in these seeds utilize during the germination of seed and their cotyledons are thin
and membranous e.g. Most of Monocot seeds e.g. Wheat, Rice, and Maize etc.
Endospermic dicot seeds : e.g., Castor, Papaya, Cotton.
Endospermic monocot seeds : e.g., Maize, Rice, Wheat and Coconut.



Bean seed is kidney shaped having a concave side and a convex side. The concave side is slightly darker and
posses a white elongated scar is called hilum, which marks the point of attachment of the seed with the
funiculus. A small pore lies adjacent to the hilum, called micropyle, which performs the function of absorbing
water during germination. There is a short ridge on the other side of hilum. It is called raphe.
The Bean seed has two seed coats. The outer seed coat is called testa. It is hard, thick, smooth and variously
coloured. It develops from outer integument of ovule. The inner coat is white, membranous and thin. It is called
tegmen. It is difficult to separate tegmen from testa. The seed coats can easily be removed if the seeds are
soaked in water for a few hours.

E 37

Micropyle Micropyle Radicle Radicle

Seed coat Radicle Plumule Cotyledon

Hilum Micropyle Radicle Seed

A coat B C D


Caruncle Radicle
Seed coat Cotyledons Plumule Cotyledon Cotyledon

Radicle Hilum
Radicle Seed
A B C D coat C


Each seed encloses an embryo which can be examined after removing the seed coats. The bulk of embryo
consists of two large, white and kidney-shaped cotyledons (or seed leaves). They store reserve food material
in the form of carbohydrates and proteins to provide nourishment to embryo at the time of seed germination
and growth of young seedling. The cotyledons are attached laterally to the curved embryonal axis at the
cotyledonary node. The micropylar end of embryonal axis is rod-shaped and slightly bulged. It is called
radicle (or the embryonic root). The other side of embryonal axis lies in between the two cotyledons. It is called
plumule (or the shoot tip). The shoot tip remains protected by two small, folded leaves. The portion of embryo
axis between the plumule and cotyledonary node is called epicotyl. The portion of axis between the radicle and
cotyledonary node is called hypocotyl.


A grain of Maize (Zea mays) is infect a dry one-seeded indehiscent fruit of caryopsis type in which pericarp
(or fruit wall) is fused with the seed coat. The covering of grain encloses two major parts-endosperm and an
embryo. As seen in longitudinal section, the major portion of grain is occupied by a large starchy endosperm.
It is surrounded by a special one-celled thick layer, called aleurone layer. The cells of aleurone layer are
filled with proteineous grains which play an important role at the time of germination.

The embryo consists of an upper shield-shaped scutellum which is actually a cotyledon. The scutellum is closely
pressed against the endosperm and helps in the translocation of nutrients from endosperm to the growing
embryo at the time of germination and seed growth. The epidermal tissue of scutellum lying in direct contact
with the endosperm is both secretory and absorptive in function. It secretes certain hormones into the endosperm
which synthesize enzymes. These enzymes are responsible for the decomposition of stored nutrients and convert
them into soluble state. The soluble nutrients are then absorbed by epithelial layer of scutellum and transferred

to the growing embryo.

The scutellum is attached laterally to the embryonal axis. One end of the embryonal axis points towards the
pointed end of the grain. It is called radicle. The radicle is covered by a protective sheath called coleorhiza.
The other end of embryonal axis, which faces towards the broader end of the grain, is called plumule. The
plumule is surrounded by a conical protective sheath, called coleoptile. The coleoptile is capable in growth and
covers the growing shoot tip till it passes through the soil during germination.

38 E

Seed coat & Pericarp Aleurone

Remnants layer
of style

Starchy endosperm


Scutellum (cotyledon) Plumule

Spathe B



The process by which the dormant embryo wakes up [becomes active] and begins to grow is known as germination
of seed. The embryo of seed grows in to a sporophyte plant. The radicle produces primary root. The plumule
develops short and latter on cotyledons degenerates.
Types of seed Germination :
Germination is basically of two types, depending upon behaviour of cotyledons but viviparous germination also
found :-
1. Epigeal germination :
Here due to hypocotyl growth or elongation, cotyledons are pushed out of soil. This is called epigeal germination.
This type of germination occurs in Cotton, Papaya, Castor, Onion, Cucurbits, Tamarind, French bean,
mustard, etc. In some cases, these above ground cotyledons become green leaf like (cotyledonary leaves)
and perform photosynthetic function till the seedling assumes independency (e.g., castor, cotton, onion, papaya,
etc.). In other plants, cotyledons do not assume leaf like shape and fall off (e.g., French bean and Tamarind).
2. Hypogeal germination :
Here due to growth in epicotyl plumule comes out of the ground and cotyledons remain underground. This is called
hypogeal germination. This type of germination occurs in most of the monocytyledons and few dicotyledons,
e.g., Maize, Rice, Wheat, Coconut, Gram, Pea, Peanut and Mango, etc.

Growing point

First leaf
Foliage leaf Broken testa
First leaf
Radicle Curved Straight Green
hypocotyl hypocotyl cotyledon
Hypocotyl Soil

Seed coat Primary root

Radicle testa Secondary
Primary root



E 39


Plumule Plumile
sheath Cotyledon

t s us
roo titio

Primary root Root hairs

(a) (b) (c) (d) (e)

(a) (b) (c) (d) (e)


Young shoot Hilum Radicle

Seed coat Strophiole Plumule


Primary root Cotyledon Radicle

Secondary roots



3. Vivipary :

It is a special type of seed germination

which is characteristic of Mangrove
vegetation, found in muddy, saline
conditions, e.g., Rhi z op hor a ,
Avicennia, Ceriops, Breguira, Branch
Sonneratia, etc. Here there is no resting
period of embryo and germination occurs

inside the fruit, while it is attached to the Calyx

parent plant, i.e., "in-situ germination". Fruit
The seedling is separated in the mud with soil
the help of lateral roots developing from
Hypocotyl Lateral roots
basal end of radicle. This is called
viviparous germination or vivipary.

40 E

The seed germination takes place in the presence of favourable condition such as temperature, moisture,
and air. This is a very delicated stage in the life of the plant. Any minute change in the environment have
harmful effect on growing embryo. So favourable environment should be essential for germination. The
following factors are essential for germination of seed.

1. Moisture :

The moisture or water is the most important factor for germination of seed. Generally, the cells of embryo
contain about 10-15% water in the dormancy period. The vital activities like growth and development is
unable to continue in this less amount of water. For active life processes, water must be present about 75-
90%. The seed absorbs water and swell up to increase in size before the germination. Water absorbs through
seed coat and micropyle. The following effects have been observed in seed due to absorption of water -

I. The seed coat becomes soft and absorbed water reaches to the cells of embryo. When the ratio of water
increases in the cells, the protoplasmic contents become more active.

II. The size of embryo increased due to water absorption due to this, seed coat breaks up and radicle and
plumule easily comes out. Seed coat absorbs water, it becomes soft and increase its permeability. Resulting
it increases rate of exchange of oxygen and CO2. The rate of respiration increases in active cells of
embryo. Thus it requires more oxygen.

III. The enzymes are present in the cells of endosperm or cotyledons which are active only in the presence of
water. The stored food changed in to dissolved form by the activity of enzymes and it reaches to the
active embryo.

2. Oxygen [O2]

The process like cell division, cell elongation etc. of the embryo require energy. This energy is released by
the oxidation of organic substances. Oxygen is essential for oxidation process. The upper surface of soil contains
sufficient amount of O2. The healthy germination does not take place in the absence or scarcity of oxygen in
deep soil so crop seed sown in the soil by the farmers upto 5-7 c.m. deep.

3. Temperature :

The suitable temperature is essential for germination of seed. The protoplasm of the cell remains active at
certain range of temperature. Most of seed do not germinate in between the range of 00C to 50C and above
the 450C. The favourable range of temperature is 20-250C for germination of seed.

4. Food or Nutrition :

The growing embryo requires [needs] nutrition during germination. The embryo depends upon stored food
materials in cotyledons or endosperm in the germination period upto the formation of primary root from the
radicle and first leaf from the plumule.

5. Light :

Light produces different effect in different variety of seeds. Most of the plants do not require light up to the
formation of first leaf. The rate of germination very fast in the absence of light. But for some plants light is very
essential for germination. They will not germinate in the dark. For example seed of Orchids, Tobacco,
Mistletoe etc.

After the development of newly leaves on shoot light becomes very essential factor. The chlorophyll is not
formed in the absence of light and this new formed plant [juvenile] will die in the absence of photosynthesis and
deficiency of food.

E 41
1. Highest amount of fat is found in endosperm of Coconut.
2. Embryonic development of Capsella is endoscopic because it is developed towards chalazal region of the
3. Single celled suspensor is found in Triticum [ wheat] and Caryophyllum. Their seeds are caryopsis fruit.
4. Suspensor is absent in Sunflower and Marigold.
5. 125 meiotic divisions are essential for development of 100 grains of Wheat.
6. Two generation and three genotypic cells are present in Angiospermic seed.
7. Heaviest seed (6 k.g.) is found in Lodoshia. Its fruit is 1 meter in length and wt. of fruit is 18kg.
8. Edible part of Betelnut is endosperm.
9. Abscisic acid is the natural germination inhibitor of seed.
10. Anatomical transition from root to stem occurs in hypocotyl region.
11. Seed of the Ruella dispersed by jaculator technique.
12. Smallest or minute seeds are found in Orchids which are lightest in plant kingdom and are called "Dust
seeds" [wt. 20.33 g].
13. Xenia (by Focke) - Effect of pollens inside Embryosac on Endosperm (except Embryo) e.g Maize.
14. Metaxenia - Effect of pollen out side the Embryosac on seed coat or pericarp. In Datepalm, maturity time as
well as size of fruits can be changed by using different pollens.
15. The pollen grains of insect pollinated flower bears oily layer is called pollen-kitt. It is made up of lipid and
16. The longest pollen tube is found in Maize.
17. Seeds of a large number of species live for several years. Some seeds can remain alive for hunderds
of years. There are several records of very old yet viable seeds. The oldest is that of a lupine,
Lupinus arcticus excavated from Arctic Tundra. The seed germinated and flowered after an esti-
mated record of 10,000 years of dormancy. A recent record of 2000 years old viable seed is of the
date plam, Phoenix dactylifera discovered during the archeological excavation at King Herod's
palace near the Dead Sea.
18. The endosperm is formed after the fertilization in Angiosperms. It is triploid while in Gymnosperm, en-
dosperm is formed before the fertilization and haploid in nature.
19. Normally, Dicotyledon embryonic development is Crucifer-type which is also known as onagrad-types.
20. The ovules of some parasitic plants do not have integument e.g. Sandal [Santalum] and Viscum.
21. The polysiphonous condition is found in families of Malvaceae and cucurbitaceae.
22. Those plants which live immortal through the seeds are known as therophytes.
23. The synergids are absent in the embryosac of Plumbago.
24. The pollination takes place at two celled structure of pollen grains in most of Angiosperms.

25. Monosporic embryosac extensively found in Angiosperm. It is discovered by Strasburger in Polygonum

divarictum. It develops only from one functional megaspore.
26. The seed of Cuscuta and Santalum lacks of cotyledon
27. The middle layer is absent in Wolffia
28. Costum and Nicodia plants have multilayered tapetum.
29. The dormancy is absent in the seed of most of Crop plants and Mangroove plants like Rhizophora.

42 E
30. The fruit formed from the ovary without fertilization is called "Parthenocarpic fruits".

31. Lotus, Nymphaea and Alisma like aquatic plants are entomophilous.

32. All the microspores of anther of Calotropis are joined together to form a special, structure called Pollinium.
These staminal structures are called translator apparatus.

33. In some plants like Typha, Drosera, Cryptostegia all four microspores joined together and they are called
Compound pollen.

34. For the formation of mature pollen grain, One meiotic and one mitotic divisions required. For the formatation of
mature male gametophyte, one meiotic and two mitotic divisions required.

35. In orchis maculata and Rumex, T-shaped tetrad of megaspore is formed (In most angiosperms generally
linear letrads are formed. Isobilateral, tetrahedral and decussate type tetrad of megaspores are absent).

36. Balanophora and Oenothera plants one micropylar megaspore of tetrad becomes functional rest of three
which are present towards the chalaza degenerate.

37. When entry of male gametes into the embryosac are not participate in fusion is called Semigamy.

38. When two pollen tube enter into an ovule and release their contents. It is possible to that the one male gamete
of one pollen tube fertilize with egg cell and one male gamete of another pollen tube participate in triple fusion
(with secondary nucleus). It is called heterofertilization. Because of this xenia effect found in Zea mays.

39. The white fluid is found in green coconut is nuclear endosperm which is known as "liquid syncytiumonlyonlyonly

40. In Cereals [Monocots] outer layer of endosperm is thick walled with dense cytoplasm, which is mainly filled with
aleurone grains [highly protein rich] and hence called Aleurone Layer. This layer secrete some hydrolytic
enzymes like proteases and amylases.


The presence of dormancy in a seed is the most important characteristic feature of plants. Because of this
character, seeds remain viable for many years. The seeds are dispersed very far places through water, air or
insects. Most of the seeds are unable to germinate just after dispersal. They germinate after sometime. The time
between the maturation and germination of seed is known as "Dormancy period". The state of inhibited
germination as a result of internal causes is usually called 'dormancy'. This seed dormancy is of considerable
advantage to the plant which helps to pass on adverse environmental conditions. The embryo remains inactive
in this period and all the growth processes suspended temporarily. There are three main basic reasons of
dormancy of seed :-


1. Impermeability of seed coat

The seed coats of many species of Leguminosae and Convolulaceae families are completely impermeable
to moisture [water] and oxygen at the time of their maturity. Seed coats, in these are thick and hard. Their
cell wall is covered by a layer of lignin which is water proof coating. Such seeds take more time for germination.
Under natural conditions seed dormancy is gradually over come by the action of microbes [bacteria] in soil, in
the alimentary canal of fruit eater birds and due to the presence of low or high temperature.

E 43
2. Dormant Embryo
In many species, although, the embryo completely not matured when the seed is ripe, even than it fails to
germinates when ever the environmental conditions are favourable or even seed coat is removed. This is
known as "Embryo" dormancy". Normally embryo dormancy occurs in many fruits yielding plants of
forest. This condition is achieved due to physiological action of seed. Such type of seed must complete their
enzymatic and chemical reaction before the germination of seed. In the lack of these reactions, seeds are
unable to germinate. such seed are kept in low temperature and in desirable moisture. Due to this seed
dormancy can be broken. The low temperature is the main reason for germination of seeds of Apple, Peaches,
Pears, Mapple and Pine etc.

3. Germination Inhibitors

Germination of some seeds is sometimes checked or prevented by the presence of some chemical compounds
are called "Germination Inhibitors" such as Ferulic acid present in Tomato juice, Caumarin, Abscisic
Acid, Dormin and Para ascorbic acid etc. This condition is generally found in xerophytic plants. These
germination inhibitors washed away with water is known as "Leaching".


(i) Scarification :- The hard seed coat is broken in this method so that water and oxygen enter into the
seed coat. The dormancy of seeds can be broken by different artificial methods -
(i) By making minute hole/pores with help of pointed sharp apparatus on the seed coat.
(ii) The seeds are rubbed on hard object so that seed coat become thin.
(iii) The partial degeneration of seed coat is carried by the action of sulphuric acid.
(ii) Stratification :- The seed kept at low temperature and in the presence of oxygen and water for
some time, so that embryo can completes its maturation.
(iii) Light Requirement :- The seeds which are affected by the light are known as "Photoblastic seeds".
The seeds which germinate in the presence of light are called "positive photoblastic seeds", such as
Lettuce, Capsella, Lepidium and Tobacco etc. The seeds in which germination take place in the
absence of light are called "negative photoblastic seeds", such as Nigella and Silene etc. Some
seeds are not affected by light are called "nonphotoblastic seeds".
The dormancy of photoblastic seeds can be broken by the treatment of red light. The phytochrome
red, absorbs red light and convert into phytochrome far red which increase the germination of seed.
Seed + R [red light] Germination
Seed + R + FR No germination
Seed + R + FR + R Germination
Seed + R + FR + R + FR No germination

(iv) Alternative Temperature :- Seeds are treated with high and low temperature alternatively at high
pressure to increase the germination of seed. e.g.The seed of Malilotus (Sweet clover) and Alfa - alfa
are treated at 2000 atmospheric pressure and 180C temperature.
(v) Viability :- This is called existance of life in a seed. The viability of seed can be tested out by 2, 3, 5,
triphenyl tetrazolium chloride. The embryonal axis of living seed becomes pink in colour in the
solution of T.T.C.
(vi) Seed dormancy can also be broken by the treatment of G.A.

44 E
(1) Hard and impermeable seed coat is found in most of legume plants which is the main reason of seed
dormancy. A hard lignin water resistant layer [seed coat] is present on seed of some Leguminoseae plants.
(2) Growth inhibitor is present in the endosperm of Iris, in seed coat of Cucurbita and in the embryo of
Xanthium .
(3) The seed dormancy in Eranthis heemalis and Ginkgo is present due to immature embryo.
(4) Some chemicals have been used successfully to interrupt seed dormancy caused by resistant or hard
impermeable seed coat. such as potassium nitrate, Ethylene, Chlorohydrine and Thiourea etc.
(5) Atriplex seed contains high concentration of salt of the time of dispersal, which do not allow germination.


Before showing their respective growth, the bud of many plants undergoes a dormant phase. This is known as
bud dormancy. This period starts in late summer and terminate in the spring season.
According to Hemberg in woody plant, the bud dormancy is caused by Abscisic acid. The level of endogenous
ABA increases with the onset of dormant period and decreases when it is broken. In non woody plant like
Potato, the bud dormancy is again due to the presence of an inhibitor. Bannet Clark and Kefford (1953)
identified this substance as Inhibitor-. It present in the peel of dormant Potato tuber. The inhibitor-is
responsible for checking the sprouting of buds located in the 'eyes' of Potato tuber.
According to Wareing. The bud dormancy, at least in woody plant, is caused by short day length, while it is
broken by long day.


(i) Chilling : The bud dormancy of some plant can be broken, if they are given cold temperature treatment
for specific duration.
(ii) Alternating temperature treatment : The dormancy of buds of some plants can be broken, if they are
subjected to low temperature (0-10C) for a brief duration and then given warm treatment.

(iii) High temperature treatment : If dry Potato tubers are stored at 35C or moist at 20C, the dormancy of
tubers buds is broken.
(iv) Chemicals : Some chemicals like 2-chloro ethanol, Gibberellin and Thiourea are capable of breaking of
dormancy of buds. Of these 2-chloro ehtanol is very effective in breaking the dormancy of Potato tuber's
buds. Endogenous Gibberellin plays a very significant role in controlling the dormancy of Potato tubers.

E 45