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SEXUAL REPRODUCTION
IN FLOWERING PLANTS
AJAY KUMAR GAUTAM
CLASS - XII
CHAPTER - 2
Plant Embryology
 The science of the origin and formation of new plants is
known as plant embryology.
 In a broader sense, plant embryology deals with the study of
all events starting from microsporogenesis,
megasporogenesis, pollination and fertilization till the
development of mature embryo.
Structure of Flower
 Flower is highly “modified condensed shoot”. It is the most conspicuous part of
angiosperms. A typical flower consists of following parts:
 Pedicel: It is a stalk or basal portion of the flower.
 Thalamus: It is also known as receptacle. It is upper swollen part of pedicel.
 Floral whorls: When the floral leaves are arranged on thalamus in cyclic
manner, they are called floral whorls. A complete flower shows the presence of
four floral whorls i.e.
1. Calyx- Individual member is sepal.
2. Corolla- Individual member is petal.
3. Androecium- Individual member is Stamen.
4. Gynoecium- Individual member is Pistil/ Carpel.
 Androecium is the male sex organ and Gynoecium is the
female sex organ.
 Each stamen is differentiated into Anther, Connective and
Filament.
 Each carpel is differentiated into Stigma, Style and Ovary.
 Presence of both androecium and gynoecium in a flower is
known as bisexual or hermaphrodite flower.
 Absence of any one sex in a flower is known as unisexual
flower.
Structure of Flower
Structure of Flower
PRE – FERTILIZATION :
STRUCTURE AND EVENTS
 Figure shows the two parts of a typical stamen - the long and slender
stalk called the filament, and the terminal generally bilobed structure
called the anther. The proximal end of the filament is attached to the
thalamus or the petal of the flower.
 The number and length of stamens are variable in flowers of different
species.
 A typical angiosperm anther is bilobed with each lobe having two
theca, i.e., they are dithecous.
 The bilobed nature of an anther is very distinct in the transverse section
of the anther. The anther is a four-sided(tetragonal) structure consisting
of four microsporangia located at the corners, two in each love.
 The microsporangia develop further and become pollen sacs. They
extend longitudinally all through the length of an anther and are packed
with pollen grains.
Stamen, Microsporangium and Pollen Grains
 Structure of microsporangium: In a transverse section, a typical
microsporangium appears near circular in outline.
 It is generally surrounded by four wall layers -the Epidermis,
Endothecium, Middle layers and the Tapetum.
 The outer three wall layers perform the function of protection and
help in dehiscence of anther to release the pollen.
 The innermost wall layer is the tapetum. It nourishes the
developing pollen grains.
 Cells of the tapetum possess dense cytoplasm and generally have
more than one nucleus.
 When the anther is young, a group of compactly arranged
homogenous cells called the sporogenous tissue occupies the
centre of each microsporangium.
Stamen, Microsporangium and Pollen Grains
 Microsporogenesis: As the anther develops, the cells of the
sporogenous tissue undergo meiotic divisions to form microspore
tetrads.
 As each cell of the sporogenous tissue is capable of giving rise to a
microspore tetrad. Each one is a potential pollen or microspore mother
cell.
 The process of formation of microspores from a pollen mother cell
(PMC) through meiosis is called microsporogenesis. The microspores,
as they are formed, are arranged in a cluster of four cells-the
microspore tetrad.
 As the anthers mature and dehydrate, the microspores dissociate from
each other and develop into pollen grains .
 Inside each microsporangium several thousands of microspores or
pollen grains are formed that are released with the dehiscence of
anther.
Stamen , Microsporangium and Pollen Grains
 Pollen grain: The pollen grains represent the male gametophytes.
 Pollen grains are generally spherical measuring about 25-50 micrometers in diameter.
 It has a prominent two-layered wall. The hard outer layer called the exine is made up
of Sporopollenin which is one of the most resistant organic material known.
 It can withstand high temperatures and strong acids and alkali. No enzyme that
degrades sporopollenin is so far known.
 Pollen grain exine has prominent apertures called germ pores where sporopollenin is
absent. Pollen grains are well-preserved as fossils because of the presence of
sporopollenin.
 The inner wall of the pollen grain is called the intine. It is a thin and continuous layer
made up of cellulose and pectin.
 The cytoplasm of pollen grain is surrounded by a plasma membrane. When the pollen
grain is mature it contains two cells, the Vegetative cell and Generative cell.
 The vegetative cell is bigger, has abundant food reserve and a large irregularly shaped
nucleus. The generative cell is small and floats in the cytoplasm of the vegetative cell
. It is spindle shaped with dense cytoplasm and a nucleus.
Stamen , Microsporangium and Pollen Grains
 Pollen grains of many species cause severe allergies and
bronchial aitlictions in some people often leading to
chronic respiratory aisorders- asthma, bronchitis, etc.
 It may be mentioned that Parthenium or carrot grass that
came into India as a contaminant with imported wheat, has
become ubiquitous in occurrence and causes pollen allergy.
 Pollen grains are rich in nutrients. It has become a fashion
in recent years to use pollen tablets 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.
Stamen , Microsporangium and Pollen Grains
The Pistil, Megasporangium (Ovule) and
Embryo Sac
 The gynoecium represents the female reproductive part of the flower.
 The gynoecium may consist of a single pistil (monocarpellary) or may nave
more than one pistil (multicarpellary).
 When there are more than one, the pistils may be fused together (syncarpous) or
may be free (apocarpous).
 Each pistil has three parts, the stigma, style and ovary.
 The stigma serves as a landing platform for pollen grains. The style is the
elongated slender part beneath the stigma. The basal bulged part of the pistil is the
ovary. Inside the ovary is the ovarian cavity (locule).The placenta is located inside
the ovarian cavity.
 Arising from the placenta are the megasporangia, commonly called ovules. The
number of Ovules in an ovary may be one (wheat, paddy, mango) to many
(papaya, water melon, orchids).
The Pistil, Megasporangium (Ovule) and
Embryo Sac
B) Multicarpellary, syncarpous pistil of
Papaver
A) A dissected flower of Hibiscus
showing pistil. ( Monocarpous)
C) A Multicarpellary, Apocarpous
Gynoecium of Michelia
 The Megasporangium (Ovule) : The ovule is a small structure attached to
the placenta by means of a stalk called funicle . The body of the ovule fuses with
funicle in the region called hilum.
 Thus, hilum represents the junction between ovule and funicle. Each ovule has one
or two protective envelopes called integuments . Integuments encircle the nucellus
except at the tip where a small opening called the micropyle is organised. Opposite
the micropylar end, is the chalaza, representing the basal part of the ovule.
 Enclosed within the integuments is a mass of cells called the nucellus. Cells of the
nucellus have abundant reserve food materials. Located in the nucellus is the
embryo sac or female gametophyte. An ovule generally has a single embryo sac
formed from a megaspore.
The Pistil, Megasporangium (Ovule) and
Embryo Sac
 Megasporogenesis : The process of formation of megaspores from the megaspore mother cell is
called megasporogenesis. Ovules generally differentiate a single megaspore mother cell (MMC) in the
micropylar region or the nucellus . It is a large cell containing dense cytoplasm and a prominent nucleus. The
MMC undergoes meiotic division. Meiosis results in the production of four megaspores.
 Female gametophyte : In a majority of flowering plants, one of the megaspores is functional while the
other three degenerate. Only the functional megaspore develops into the female gametophyte(embryo sac).
This method of embryo sac formation from a single megaspore is termed monosporic development.
 The nucleus of the functional megaspore divides mitotically to form two nuclei which move to the opposite
poles, forming the 2-nucleate embryo sac. Two more sequential mitotic nuclear divisions result in the
formation of the 4-nucleate and later the 8-nucleate stages of the embryo sac.
 After the 8-nucleate stage, cell walls are laid down leading to the organisation of the typical female
gametophyte or embryo sac. Observe the distribution of cells inside the embryo sac. Six of the eight nuclei
are surrounded by cell walls and organised into cells; the remaining two nuclei, called polar nuclei are
situated below the egg apparatus in the large central cell.
The Pistil, Megasporangium (Ovule) and
Embryo Sac
 There is a characteristic distribution of the cells within the embryo sac. Three cells are grouped
together at the micropylar end and constitute the egg apparatus. The egg apparatus, in turn,
consists of two synergids and one egg cell.
 The synergids have special cellular thickenings at the micropylar tip called filiform
apparatus, which play an important role in guiding the pollen tubes into the synergid.
 Three cells are at the chalazal end and are called the antipodals . The large central cell, as
mentioned earlier, has two polar nuclei. Thus, a typical angiosperm embryo sac, at maturity,
though 8-nucleate is 7-celled.
The Pistil, Megasporangium (Ovule) and
Embryo Sac
POLLINATION
 Pollination is the mechanism to achieve the objective of transfer of pollen grains (shed from
the anther) to the stigma of a pistil is termed pollination. Flowering plants have evolved an
amazing array of adaptations to achieve pollination. They make use of external agents to
achieve pollination.
 Kinds of Pollination : Depending on the source of pollen, pollination can be divided into three
types.
AUTOGAMY GEITONOGAMY
XENOGAMY
POLLINATION
I. Autogamy: The transfer of pollen grain from the anther to the stigma of the same flower.
 Adaptations seen in plant to ensure self-pollination :
1. Bisexuality : Presence of both the essential whorls in the same flower.
2. Homogamy : Maturation of both androecium and gynoecium at the same time, i.e., there should be
synchrony in release of pollen and stigma maturation.
3. Cleistogamy : A condition in which flower does not open. In such flowers, the anthers and stigma
lie close to each other. When anther dehisce in the flower buds, pollen grains come in contact with
stigma to effect pollination.
 Some plants like Viola (common pansy), Oxalis and Commelina produce both types of flowers, i.e.,
Open Flowers (Chasmogamous) and Closed Flowers ( Cleistogamous).
TYPES OF POLLINATION
 Advantage of Cleistogamy :
1. It ensures seed formation even in the absence of any pollinating agent.
2. It is cheaper for the plant as there is no costly nectar or fragrance which the plant has to produce for
pollination.
 Disadvantage of Cleistogamy: The offsprings produced have limited genetic diversity.
1. Bud pollination : When self pollination occurs in the bud stage before the opening of flowers e.g.
Pea, Rice.
II. Geitnogamy : The transfer of pollen grains from anther to the stigma of another
flower of the same plant. This transfer involves an agent of pollination, hence functionally it is
cross-pollination. Genetically, it Is similar to autogamy since the pollen grains come from the
same plant.
TYPES OF POLLINATION
III. Xenogamy : The transfer of pollen grains from anther to the stigma of
another flower of different plant of the same species.
 This is the only type of pollination which brings genetically different types of pollen
grains to stigma.
Abiotic Agents
 Wind (Anemophily): This is the more common amongst abiotic pollinations.
 Characteristics of wind pollinated flowers :
1. The pollen grains are light and non-sticky, so that they can easily be carried by air
currents.
2. The flowers have well-exposed stamens, so that the pollens are easily dispersed into
wind currents.
3. Flowers have large feathery stigma to easily trap the air borne pollen grains.
4. Nectaries absent.
5. Presence of single ovule in each ovary.
6. Flowers packed into inflorescence.
TYPES OF POLLINATION
Abiotic Agents
 Examples of wind pollinated plants: It is quite common in grasses.
a) Maize
b) Wheat
c) Sugarcane
d) Bamboo
 Water (Hydrophily): Pollination by water is quite rare in flowering plants and is limited to about 30 genera, mostly
monocots, e.g., Fresh water plants like Vallisneria , Hydrilla ; marine water plant like Zoostera.
 Characteristics Features:
I. Light, unwettable pollen grains. Generally, surrounded by mucilagenous covering, hence protected from wetting.
II. Long, sticky wettable stigma.
Pollination by water may occur at two places ;
i. On the surface of water ( Epihydrophily) : Example : Vallisneria
 the female flower have a very long pedicel, therefore it reaches the surface of water.
 Male flowers after breakage floats on the surface of water.
 Pollen grains are released on to the surface of water. They are carried passively by water currents, some of them eventually reach
the female flowers and the stigma
 Beneath the surface of water ( Hypohydrophily ): Example : Zoostera (sea grass)
 Zoostera is a marine water plant. Female flowers remain submerged in water. Pollen grains are long
ribbon like and they are carried passively inside the water, some of them reach the stigma and
achieve pollination.
 In a majority of aquatic plants, the flowers emerge above the level of water and are pollinated by
insects or wind. E.g., Water hyacinth and water lily are pollinated by insects.
 Flowers pollinated by abiotic agents are not very colourful and do not produce nectar because the
bright colours of flowers are to attract insects to pollinate them and the nectar they give is a reward
for the pollinator. Pollen grains coming in contact with stigma is a chance factor in both wind and
water pollination. To compensate for these uncertainties and associated loss of pollen grains, the
flowers produce enormous amount of pollen when compared to the number of ovules available for
pollination.
Abiotic Agents
 Majority of flowering plants use a range of animals as pollinating agents. Bees, butterflies, flies, beetles, wasps,
ants, moths, birds (sun birds and humming birds) and bats are the common pollinating agents.
 Larger animals like some primates (Lemur), arboreal (tree- dwelling ) rodents, or even reptiles (gecko lizard and
garden lizard) have also been reported as pollinators in some species. E.g., Lemur in Ravenela plant and lizard in
flax.
 Insects ( Entomophily) : These are the most common biotic agents of pollination. Bees are the most
common insect which acts as a pollinating agents. Other insect pollinators are butterflies, flies, beetles, wasps,
ants, moths.
 Characteristic features of flowers pollinated by insects:
 Majority of insect-pollinated flowers are large-sized.
 Small-sized flowers are clustered into an inflorescence.
 Colourful, fragrant.
 Nectaries present.
 Sticky pollen grain.
Biotic Agents
 Floral rewards for insects:
 Nectar
 Pollen
 Safe place to lay eggs. For example, Amorphophallus (flower is 6 feet in height).
 Pronuba moth lays its eggs in the ovary of Yucca plant and its flowers get pollinated by the moth. Both the
organisms cannot complete their lifecycle without each other. The moth deposits its eggs in the locule of the
ovary and the flower, in turn, gets pollinated by the moth. The larvae of the moth come out of the eggs as seeds
start developing.
 Pollen/nectar Robbers Insects which consume pollen or nectar without bringing about pollination, e.g., Queens
of Bombus affinis perforate Aquilegia spurs and steal nectar.
 Examples of Insect Pollinated Flowers Yucca, Salvia, Ficus , Calotropis , Centaurea , Aristolochia .
Biotic Agents
Outbreeding Devices
 Outbreeding Devices
 Devices or features of plants which discourage self-pollination and encourage cross-pollination are called as
outbreeding devices. It prevents the loss in genetic variation or inbreeding depression which may be a result of
continued self-pollination .
 Examples of these devices:
 Unisexuality : Formation of unisexual flowers. Two conditions are their.
(a) Monoecious plants : Male and female flowers are produced on the same plant. This prevents autogamy but not
geitnogamy. E.g., Castor, maize.
(b) Dioecious : Male and female flowers are produced on two separate plants. T prevents autogamy as well as
geitnogamy. E.g., Vallisneria, Papaya.
(c) Dichogamy : Different maturation time of androecium and gynoecium in the same flower. Either the pollen is
released before the stigma becomes receptive ( protandrous, e.g., Sunflower, Cotton) or stigma becomes receptive
much before the release of pollen ( protogynous, e.g., Ficus, Aristolochia ).
(d) Self-incompatibility : The failure of the pollen grains (from the same flower or other flowers of the same plant)
from fertilising the ovule by inhibiting pollen germination or pollen tube growth in the pistil. It is a genetically
controlled mechanism.
 In some species, the anther and stigma are placed at different position, so that pollen cannot come in contact with the
stigma of the same flower.
Pollen-Pistil Interaction
a) Pollination Does not guarantee the transfer of the right type of pollen on stigma.
Often, pollen of the wrong type, either from other species or from the same plant (if it
is self-incompatible), also land on the stigma.
b) The pistil has the ability to recognise the pollen, whether it is of the right type
(compatible) or of the wrong type (incompatible).
c) The ability of the pistil to recognise the pollen followed by its acceptance or rejection
is the result of a continuous chemical dialogue between pollen grain and the pistil.
Post-pollination events after the compatible pollination are listed
below :
1. The pollen grain germinates on the stigma to produce a pollen tube through one of the
germ pores. The contents of the pollen grain move into the pollen tube.
2. Pollen tube grows through the tissues of the stigma and style and reaches ovary.
Growth of pollen tube is chemotropic.
3. The generative cell divides and forms two male gametes during the growth of the
pollen tube in the stigma, if pollen grains are shed at two-celled condition.
4. If pollen grains are shed at three-celled stage, pollen tube carries two male gametes
from the beginning.
 Entry of pollen tube into embryo sac : Irrespective of the place
of entry of pollen tube into ovule, the tube invariably enters embryo sac at
micropylar end, i.e., degenerating synergid cell.
• Many recent studies have shown that filiform apparatus present at
micropylar part of synergids guides the entry of pollen tube.
• All these events from pollen deposition on the stigma until pollen tube
enters the ovule are together referred as pollen-pistil interaction.
Pollen-Pistil Interaction
Entry of pollen tube into ovule
takes place through
POROGAMY CHALAZOGAMY MESOGAMY
This is the most common e.g. Casuarina e.g., Cucurbita
condition and takes place
in most of the flowering
plants.
 In-Vitro pollen germination: Pollen germination can be studied by dusting pollen (e.g., pea, Chick pea, Crotalaria, balsam,
Vinca) on a glass slide containing a drop of 10% sugar solution with boric acid, Ca, Mg and K salts. After 15-30 minutes, pollen tubes
will be observed to come out of the pollen grains. So, this germination of pollen grain in laboratory is called Hanging drop method.
Let us try to know how the pollination can be altered to obtain the superior varieties.
 Artificial hybridisation: It is a method of crop improvement in which crosses are made between different varieties, species and
genera, in order to combine the desirable characters in a single ‘superior variety’.
 This technique involve the following steps :
(1) Selection of suitable parents.
(2) If the female parents bears bisexual flowers in such crossing experiments, it is important to make sure that only the desired pollen grains
are used for pollination.
a) Emasculation: Removal of anthers from female parent flower buds before the anther dehisces.
b) Bagging: Covering of emasculated flowers with a bag of suitable size generally made of butter paper to prevent contamination of
stigma with unwanted pollen.
(4) Dusting of pollen grains from anthers of male parent on the stigma of female parent when stigma attains receptivity and then it is
rebagged .
(4) Fruits are then allowed to develop.
(5) If the female parent produces unisexual flowers, there is no need for emasculation.
Pollen-Pistil Interaction
Double Fertilization
 After entering one of the synergids, the pollen tube releases the two male gametes into the
cytoplasm of the synergids. The following events take place in the embryo sac :
(1) SYNGAMY :
♂Male gamete (n) + ♀Egg (n) Zygote (2n)
 Male gamete moves towards the egg cell and fuses with its nucleus. Thus, resulting in the formation
of a diploid cell, the zygote.
(2) TRIPLE FUSION :
♂Male gamete (n) + Two fused polar nuclei (2n) PEN Primary Endosperm Nucleus
 The other male gamete moves towards the two polar nuclei located in the central cell and fuses with
them to produce a triploid primary endosperm nucleus (PEN).
Post fertilization : Structures & Events
 Following double fertilization, events of endosperms and embryo development, maturation of ovule(s)
into seed(s) and ovary into fruits, are collectively termed Post Fertilization events.
 Post Fertilization events include:
I. Endosperm Development
II. Embryo development
III. Ovules maturing into seed
IV. Ovary maturing into fruit
 ENDOSPERM DEVELOPMENT :
 Endosperm development precedes embryo development. The primary endosperm cells divides
repeatedly and forms a triploid endosperm tissue. The cells of this tissue are filled with reserve food
materials and are used for the nutrition of the developing embryo.
 In the most common type of endosperm development, the PEN undergoes successive nuclear divisions
to give rise to free nuclei. This stage of endosperm development is called Free Nuclear Endosperm.
 Subsequently cell wall formation occurs and the endosperm becomes cellular. The number of free nuclei
formed before cellularisation varies greatly.
 The coconut water from green tender coconut that we are familiar with, is nothing but free nuclear
endosperm (made up of thousands of nuclei) and the surrounding white kernel is the cellular
endosperm.
Embryo Development
 Embryo develops at the micropylar end of the embryo sac where the zygote is situated. Most
zygotes divide only after certain amount of endosperm is formed. This is an adaptation to provide
assured nutrition to the developing embryo. Though the seeds differ greatly, the early stages of
embryo development (embryogeny) are similar in both monocotyledons and dicotyledons. the
stages of embryogeny in a dicotyledonous embryo.
 The zygote gives rise to the proembryo and subsequently to the globular, heart-shaped and
mature embryo.
 A typical dicotyledonous embryo consists of an embryonal axis and two cotyledons. The portion
of embryonal axis above the level of cotyledons is the epicotyl, which terminates with the
plumule or stem tip.
 The cylindrical portion below the level of cotyledons is hypocotyl that terminates at its lower end
in the radicle or root tip. The root tip is covered with a root cap.
 Embryos of monocotyledons (Figure 2.14 b) possess only one cotyledon. 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 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
coleoptile.
Seed Development
 In angiosperms, the seed is the final product of sexual reproduction. It is often described
as a fertilised ovule. Seeds are formed inside fruits. A seed typically consists of seed
coat(s), cotyledon(s) and an embryo axis.
 The cotyledons of the embryo are simple structures generally thick and swollen due to
storage of food reserves (as in legumes).
 Mature seeds may be non-albuminous or ex-albuminous :
 Non- Albuminous: Seeds have no residual endosperm as it is completely consumed
during embryo development (e.g., pea, groundnut).
 Albuminous seeds: They retain a part of endosperm as it is not completely used up
during embryo development (e.g.. Wheat, maize, barley, castor).
 Perispermic Seeds : In some seeds such as black pepper and beet, remnants of nucellus
are also persistent. This residual, persistent nucellus is the perisperm.
 Integuments of ovules harden as tough protective seed coats.
 The micropyle remains as a small pore in the seed coat. This facilitates entry of oxygen
and water into the seed during germination. As the seed matures, its water content is
reduced and seeds become relatively dry (10-15 per cent moisture by mass).
 The general metabolic activity of the embryo slows down. The embryo may enter a state
of inactivity called dormancy, or if favourable conditions are available (adequate
moisture, oxygen and suitable temperature), they germinate.
Fruit Formation
 As ovules mature into seeds, the ovary develops into a fruit, i.e., the
transformation of ovules into seeds and ovary into fruit proceeds simultaneously.
 The wall of the ovary develops into the wall of fruit called pericarp.
 The fruits may be fleshy as in guava, orange, mango, etc., or may be dry, as in
groundnut, and mustard, etc. Many fruits have evolved mechanisms for dispersal
of seeds.
 In most plants, by the time the fruit develops from the ovary, other floral parts
degenerate and fall off.
 However, in a few species such as apple, strawberry, cashew, etc., the thalamus
also contributes to fruit formation. Such fruits are called false fruits.
 Most fruits however develop only from the ovary and are called true fruits.
 Although in most of the species, fruits are the results of fertilisation, there are a
few species in which fruits develop without fertilisation. Such fruits are called
Parthenocarpic fruits. Banana is ones uchexample.
 Parthenocarpy can be induced through the application of growth hormones and
such fruits are seedless.
APOMIXIS & POLYEMBRYONY
 Although seeds, in general are the products of fertilisation, a few flowering plants such as some
species of Asteraceae and grasses. have evolved a special mechanism, to produce seeds without
fertilisation, called apomixis.
 Apomixis is a form of asexual reproduction that mimics sexual reproduction.
 In some species. The diploid egg cell is formed without reduction division and develops into the
embryo without fertilisation.
 Some of the nucellar cells surrounding the embryo sac start dividing, protrude into the embryo sac
and develop into the embryos. In such species each ovule contains many embryos. Occurrence of
more than one embryo in a seed is referred to as polyembryony.

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Sexual reproduction in flowering plants 2

  • 1. SEXUAL REPRODUCTION IN FLOWERING PLANTS AJAY KUMAR GAUTAM CLASS - XII CHAPTER - 2
  • 2. Plant Embryology  The science of the origin and formation of new plants is known as plant embryology.  In a broader sense, plant embryology deals with the study of all events starting from microsporogenesis, megasporogenesis, pollination and fertilization till the development of mature embryo.
  • 3. Structure of Flower  Flower is highly “modified condensed shoot”. It is the most conspicuous part of angiosperms. A typical flower consists of following parts:  Pedicel: It is a stalk or basal portion of the flower.  Thalamus: It is also known as receptacle. It is upper swollen part of pedicel.  Floral whorls: When the floral leaves are arranged on thalamus in cyclic manner, they are called floral whorls. A complete flower shows the presence of four floral whorls i.e. 1. Calyx- Individual member is sepal. 2. Corolla- Individual member is petal. 3. Androecium- Individual member is Stamen. 4. Gynoecium- Individual member is Pistil/ Carpel.
  • 4.  Androecium is the male sex organ and Gynoecium is the female sex organ.  Each stamen is differentiated into Anther, Connective and Filament.  Each carpel is differentiated into Stigma, Style and Ovary.  Presence of both androecium and gynoecium in a flower is known as bisexual or hermaphrodite flower.  Absence of any one sex in a flower is known as unisexual flower. Structure of Flower
  • 6. PRE – FERTILIZATION : STRUCTURE AND EVENTS
  • 7.  Figure shows the two parts of a typical stamen - the long and slender stalk called the filament, and the terminal generally bilobed structure called the anther. The proximal end of the filament is attached to the thalamus or the petal of the flower.  The number and length of stamens are variable in flowers of different species.  A typical angiosperm anther is bilobed with each lobe having two theca, i.e., they are dithecous.  The bilobed nature of an anther is very distinct in the transverse section of the anther. The anther is a four-sided(tetragonal) structure consisting of four microsporangia located at the corners, two in each love.  The microsporangia develop further and become pollen sacs. They extend longitudinally all through the length of an anther and are packed with pollen grains. Stamen, Microsporangium and Pollen Grains
  • 8.  Structure of microsporangium: In a transverse section, a typical microsporangium appears near circular in outline.  It is generally surrounded by four wall layers -the Epidermis, Endothecium, Middle layers and the Tapetum.  The outer three wall layers perform the function of protection and help in dehiscence of anther to release the pollen.  The innermost wall layer is the tapetum. It nourishes the developing pollen grains.  Cells of the tapetum possess dense cytoplasm and generally have more than one nucleus.  When the anther is young, a group of compactly arranged homogenous cells called the sporogenous tissue occupies the centre of each microsporangium. Stamen, Microsporangium and Pollen Grains
  • 9.  Microsporogenesis: As the anther develops, the cells of the sporogenous tissue undergo meiotic divisions to form microspore tetrads.  As each cell of the sporogenous tissue is capable of giving rise to a microspore tetrad. Each one is a potential pollen or microspore mother cell.  The process of formation of microspores from a pollen mother cell (PMC) through meiosis is called microsporogenesis. The microspores, as they are formed, are arranged in a cluster of four cells-the microspore tetrad.  As the anthers mature and dehydrate, the microspores dissociate from each other and develop into pollen grains .  Inside each microsporangium several thousands of microspores or pollen grains are formed that are released with the dehiscence of anther. Stamen , Microsporangium and Pollen Grains
  • 10.  Pollen grain: The pollen grains represent the male gametophytes.  Pollen grains are generally spherical measuring about 25-50 micrometers in diameter.  It has a prominent two-layered wall. The hard outer layer called the exine is made up of Sporopollenin which is one of the most resistant organic material known.  It can withstand high temperatures and strong acids and alkali. No enzyme that degrades sporopollenin is so far known.  Pollen grain exine has prominent apertures called germ pores where sporopollenin is absent. Pollen grains are well-preserved as fossils because of the presence of sporopollenin.  The inner wall of the pollen grain is called the intine. It is a thin and continuous layer made up of cellulose and pectin.  The cytoplasm of pollen grain is surrounded by a plasma membrane. When the pollen grain is mature it contains two cells, the Vegetative cell and Generative cell.  The vegetative cell is bigger, has abundant food reserve and a large irregularly shaped nucleus. The generative cell is small and floats in the cytoplasm of the vegetative cell . It is spindle shaped with dense cytoplasm and a nucleus. Stamen , Microsporangium and Pollen Grains
  • 11.  Pollen grains of many species cause severe allergies and bronchial aitlictions in some people often leading to chronic respiratory aisorders- asthma, bronchitis, etc.  It may be mentioned that Parthenium or carrot grass that came into India as a contaminant with imported wheat, has become ubiquitous in occurrence and causes pollen allergy.  Pollen grains are rich in nutrients. It has become a fashion in recent years to use pollen tablets 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. Stamen , Microsporangium and Pollen Grains
  • 12. The Pistil, Megasporangium (Ovule) and Embryo Sac  The gynoecium represents the female reproductive part of the flower.  The gynoecium may consist of a single pistil (monocarpellary) or may nave more than one pistil (multicarpellary).  When there are more than one, the pistils may be fused together (syncarpous) or may be free (apocarpous).  Each pistil has three parts, the stigma, style and ovary.  The stigma serves as a landing platform for pollen grains. The style is the elongated slender part beneath the stigma. The basal bulged part of the pistil is the ovary. Inside the ovary is the ovarian cavity (locule).The placenta is located inside the ovarian cavity.  Arising from the placenta are the megasporangia, commonly called ovules. The number of Ovules in an ovary may be one (wheat, paddy, mango) to many (papaya, water melon, orchids).
  • 13. The Pistil, Megasporangium (Ovule) and Embryo Sac B) Multicarpellary, syncarpous pistil of Papaver A) A dissected flower of Hibiscus showing pistil. ( Monocarpous) C) A Multicarpellary, Apocarpous Gynoecium of Michelia
  • 14.  The Megasporangium (Ovule) : The ovule is a small structure attached to the placenta by means of a stalk called funicle . The body of the ovule fuses with funicle in the region called hilum.  Thus, hilum represents the junction between ovule and funicle. Each ovule has one or two protective envelopes called integuments . Integuments encircle the nucellus except at the tip where a small opening called the micropyle is organised. Opposite the micropylar end, is the chalaza, representing the basal part of the ovule.  Enclosed within the integuments is a mass of cells called the nucellus. Cells of the nucellus have abundant reserve food materials. Located in the nucellus is the embryo sac or female gametophyte. An ovule generally has a single embryo sac formed from a megaspore. The Pistil, Megasporangium (Ovule) and Embryo Sac
  • 15.
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  • 17.  Megasporogenesis : The process of formation of megaspores from the megaspore mother cell is called megasporogenesis. Ovules generally differentiate a single megaspore mother cell (MMC) in the micropylar region or the nucellus . It is a large cell containing dense cytoplasm and a prominent nucleus. The MMC undergoes meiotic division. Meiosis results in the production of four megaspores.  Female gametophyte : In a majority of flowering plants, one of the megaspores is functional while the other three degenerate. Only the functional megaspore develops into the female gametophyte(embryo sac). This method of embryo sac formation from a single megaspore is termed monosporic development.  The nucleus of the functional megaspore divides mitotically to form two nuclei which move to the opposite poles, forming the 2-nucleate embryo sac. Two more sequential mitotic nuclear divisions result in the formation of the 4-nucleate and later the 8-nucleate stages of the embryo sac.  After the 8-nucleate stage, cell walls are laid down leading to the organisation of the typical female gametophyte or embryo sac. Observe the distribution of cells inside the embryo sac. Six of the eight nuclei are surrounded by cell walls and organised into cells; the remaining two nuclei, called polar nuclei are situated below the egg apparatus in the large central cell. The Pistil, Megasporangium (Ovule) and Embryo Sac
  • 18.  There is a characteristic distribution of the cells within the embryo sac. Three cells are grouped together at the micropylar end and constitute the egg apparatus. The egg apparatus, in turn, consists of two synergids and one egg cell.  The synergids have special cellular thickenings at the micropylar tip called filiform apparatus, which play an important role in guiding the pollen tubes into the synergid.  Three cells are at the chalazal end and are called the antipodals . The large central cell, as mentioned earlier, has two polar nuclei. Thus, a typical angiosperm embryo sac, at maturity, though 8-nucleate is 7-celled. The Pistil, Megasporangium (Ovule) and Embryo Sac
  • 19.
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  • 21. POLLINATION  Pollination is the mechanism to achieve the objective of transfer of pollen grains (shed from the anther) to the stigma of a pistil is termed pollination. Flowering plants have evolved an amazing array of adaptations to achieve pollination. They make use of external agents to achieve pollination.  Kinds of Pollination : Depending on the source of pollen, pollination can be divided into three types. AUTOGAMY GEITONOGAMY XENOGAMY POLLINATION
  • 22. I. Autogamy: The transfer of pollen grain from the anther to the stigma of the same flower.  Adaptations seen in plant to ensure self-pollination : 1. Bisexuality : Presence of both the essential whorls in the same flower. 2. Homogamy : Maturation of both androecium and gynoecium at the same time, i.e., there should be synchrony in release of pollen and stigma maturation. 3. Cleistogamy : A condition in which flower does not open. In such flowers, the anthers and stigma lie close to each other. When anther dehisce in the flower buds, pollen grains come in contact with stigma to effect pollination.  Some plants like Viola (common pansy), Oxalis and Commelina produce both types of flowers, i.e., Open Flowers (Chasmogamous) and Closed Flowers ( Cleistogamous). TYPES OF POLLINATION
  • 23.  Advantage of Cleistogamy : 1. It ensures seed formation even in the absence of any pollinating agent. 2. It is cheaper for the plant as there is no costly nectar or fragrance which the plant has to produce for pollination.  Disadvantage of Cleistogamy: The offsprings produced have limited genetic diversity. 1. Bud pollination : When self pollination occurs in the bud stage before the opening of flowers e.g. Pea, Rice. II. Geitnogamy : The transfer of pollen grains from anther to the stigma of another flower of the same plant. This transfer involves an agent of pollination, hence functionally it is cross-pollination. Genetically, it Is similar to autogamy since the pollen grains come from the same plant. TYPES OF POLLINATION
  • 24. III. Xenogamy : The transfer of pollen grains from anther to the stigma of another flower of different plant of the same species.  This is the only type of pollination which brings genetically different types of pollen grains to stigma. Abiotic Agents  Wind (Anemophily): This is the more common amongst abiotic pollinations.  Characteristics of wind pollinated flowers : 1. The pollen grains are light and non-sticky, so that they can easily be carried by air currents. 2. The flowers have well-exposed stamens, so that the pollens are easily dispersed into wind currents. 3. Flowers have large feathery stigma to easily trap the air borne pollen grains. 4. Nectaries absent. 5. Presence of single ovule in each ovary. 6. Flowers packed into inflorescence. TYPES OF POLLINATION
  • 25. Abiotic Agents  Examples of wind pollinated plants: It is quite common in grasses. a) Maize b) Wheat c) Sugarcane d) Bamboo  Water (Hydrophily): Pollination by water is quite rare in flowering plants and is limited to about 30 genera, mostly monocots, e.g., Fresh water plants like Vallisneria , Hydrilla ; marine water plant like Zoostera.  Characteristics Features: I. Light, unwettable pollen grains. Generally, surrounded by mucilagenous covering, hence protected from wetting. II. Long, sticky wettable stigma. Pollination by water may occur at two places ; i. On the surface of water ( Epihydrophily) : Example : Vallisneria  the female flower have a very long pedicel, therefore it reaches the surface of water.  Male flowers after breakage floats on the surface of water.  Pollen grains are released on to the surface of water. They are carried passively by water currents, some of them eventually reach the female flowers and the stigma
  • 26.  Beneath the surface of water ( Hypohydrophily ): Example : Zoostera (sea grass)  Zoostera is a marine water plant. Female flowers remain submerged in water. Pollen grains are long ribbon like and they are carried passively inside the water, some of them reach the stigma and achieve pollination.  In a majority of aquatic plants, the flowers emerge above the level of water and are pollinated by insects or wind. E.g., Water hyacinth and water lily are pollinated by insects.  Flowers pollinated by abiotic agents are not very colourful and do not produce nectar because the bright colours of flowers are to attract insects to pollinate them and the nectar they give is a reward for the pollinator. Pollen grains coming in contact with stigma is a chance factor in both wind and water pollination. To compensate for these uncertainties and associated loss of pollen grains, the flowers produce enormous amount of pollen when compared to the number of ovules available for pollination. Abiotic Agents
  • 27.  Majority of flowering plants use a range of animals as pollinating agents. Bees, butterflies, flies, beetles, wasps, ants, moths, birds (sun birds and humming birds) and bats are the common pollinating agents.  Larger animals like some primates (Lemur), arboreal (tree- dwelling ) rodents, or even reptiles (gecko lizard and garden lizard) have also been reported as pollinators in some species. E.g., Lemur in Ravenela plant and lizard in flax.  Insects ( Entomophily) : These are the most common biotic agents of pollination. Bees are the most common insect which acts as a pollinating agents. Other insect pollinators are butterflies, flies, beetles, wasps, ants, moths.  Characteristic features of flowers pollinated by insects:  Majority of insect-pollinated flowers are large-sized.  Small-sized flowers are clustered into an inflorescence.  Colourful, fragrant.  Nectaries present.  Sticky pollen grain. Biotic Agents
  • 28.  Floral rewards for insects:  Nectar  Pollen  Safe place to lay eggs. For example, Amorphophallus (flower is 6 feet in height).  Pronuba moth lays its eggs in the ovary of Yucca plant and its flowers get pollinated by the moth. Both the organisms cannot complete their lifecycle without each other. The moth deposits its eggs in the locule of the ovary and the flower, in turn, gets pollinated by the moth. The larvae of the moth come out of the eggs as seeds start developing.  Pollen/nectar Robbers Insects which consume pollen or nectar without bringing about pollination, e.g., Queens of Bombus affinis perforate Aquilegia spurs and steal nectar.  Examples of Insect Pollinated Flowers Yucca, Salvia, Ficus , Calotropis , Centaurea , Aristolochia . Biotic Agents
  • 29. Outbreeding Devices  Outbreeding Devices  Devices or features of plants which discourage self-pollination and encourage cross-pollination are called as outbreeding devices. It prevents the loss in genetic variation or inbreeding depression which may be a result of continued self-pollination .  Examples of these devices:  Unisexuality : Formation of unisexual flowers. Two conditions are their. (a) Monoecious plants : Male and female flowers are produced on the same plant. This prevents autogamy but not geitnogamy. E.g., Castor, maize. (b) Dioecious : Male and female flowers are produced on two separate plants. T prevents autogamy as well as geitnogamy. E.g., Vallisneria, Papaya. (c) Dichogamy : Different maturation time of androecium and gynoecium in the same flower. Either the pollen is released before the stigma becomes receptive ( protandrous, e.g., Sunflower, Cotton) or stigma becomes receptive much before the release of pollen ( protogynous, e.g., Ficus, Aristolochia ). (d) Self-incompatibility : The failure of the pollen grains (from the same flower or other flowers of the same plant) from fertilising the ovule by inhibiting pollen germination or pollen tube growth in the pistil. It is a genetically controlled mechanism.  In some species, the anther and stigma are placed at different position, so that pollen cannot come in contact with the stigma of the same flower.
  • 30. Pollen-Pistil Interaction a) Pollination Does not guarantee the transfer of the right type of pollen on stigma. Often, pollen of the wrong type, either from other species or from the same plant (if it is self-incompatible), also land on the stigma. b) The pistil has the ability to recognise the pollen, whether it is of the right type (compatible) or of the wrong type (incompatible). c) The ability of the pistil to recognise the pollen followed by its acceptance or rejection is the result of a continuous chemical dialogue between pollen grain and the pistil. Post-pollination events after the compatible pollination are listed below : 1. The pollen grain germinates on the stigma to produce a pollen tube through one of the germ pores. The contents of the pollen grain move into the pollen tube. 2. Pollen tube grows through the tissues of the stigma and style and reaches ovary. Growth of pollen tube is chemotropic. 3. The generative cell divides and forms two male gametes during the growth of the pollen tube in the stigma, if pollen grains are shed at two-celled condition. 4. If pollen grains are shed at three-celled stage, pollen tube carries two male gametes from the beginning.
  • 31.  Entry of pollen tube into embryo sac : Irrespective of the place of entry of pollen tube into ovule, the tube invariably enters embryo sac at micropylar end, i.e., degenerating synergid cell. • Many recent studies have shown that filiform apparatus present at micropylar part of synergids guides the entry of pollen tube. • All these events from pollen deposition on the stigma until pollen tube enters the ovule are together referred as pollen-pistil interaction. Pollen-Pistil Interaction Entry of pollen tube into ovule takes place through POROGAMY CHALAZOGAMY MESOGAMY This is the most common e.g. Casuarina e.g., Cucurbita condition and takes place in most of the flowering plants.
  • 32.  In-Vitro pollen germination: Pollen germination can be studied by dusting pollen (e.g., pea, Chick pea, Crotalaria, balsam, Vinca) on a glass slide containing a drop of 10% sugar solution with boric acid, Ca, Mg and K salts. After 15-30 minutes, pollen tubes will be observed to come out of the pollen grains. So, this germination of pollen grain in laboratory is called Hanging drop method. Let us try to know how the pollination can be altered to obtain the superior varieties.  Artificial hybridisation: It is a method of crop improvement in which crosses are made between different varieties, species and genera, in order to combine the desirable characters in a single ‘superior variety’.  This technique involve the following steps : (1) Selection of suitable parents. (2) If the female parents bears bisexual flowers in such crossing experiments, it is important to make sure that only the desired pollen grains are used for pollination. a) Emasculation: Removal of anthers from female parent flower buds before the anther dehisces. b) Bagging: Covering of emasculated flowers with a bag of suitable size generally made of butter paper to prevent contamination of stigma with unwanted pollen. (4) Dusting of pollen grains from anthers of male parent on the stigma of female parent when stigma attains receptivity and then it is rebagged . (4) Fruits are then allowed to develop. (5) If the female parent produces unisexual flowers, there is no need for emasculation. Pollen-Pistil Interaction
  • 33. Double Fertilization  After entering one of the synergids, the pollen tube releases the two male gametes into the cytoplasm of the synergids. The following events take place in the embryo sac : (1) SYNGAMY : ♂Male gamete (n) + ♀Egg (n) Zygote (2n)  Male gamete moves towards the egg cell and fuses with its nucleus. Thus, resulting in the formation of a diploid cell, the zygote. (2) TRIPLE FUSION : ♂Male gamete (n) + Two fused polar nuclei (2n) PEN Primary Endosperm Nucleus  The other male gamete moves towards the two polar nuclei located in the central cell and fuses with them to produce a triploid primary endosperm nucleus (PEN).
  • 34. Post fertilization : Structures & Events  Following double fertilization, events of endosperms and embryo development, maturation of ovule(s) into seed(s) and ovary into fruits, are collectively termed Post Fertilization events.  Post Fertilization events include: I. Endosperm Development II. Embryo development III. Ovules maturing into seed IV. Ovary maturing into fruit  ENDOSPERM DEVELOPMENT :  Endosperm development precedes embryo development. The primary endosperm cells divides repeatedly and forms a triploid endosperm tissue. The cells of this tissue are filled with reserve food materials and are used for the nutrition of the developing embryo.  In the most common type of endosperm development, the PEN undergoes successive nuclear divisions to give rise to free nuclei. This stage of endosperm development is called Free Nuclear Endosperm.  Subsequently cell wall formation occurs and the endosperm becomes cellular. The number of free nuclei formed before cellularisation varies greatly.  The coconut water from green tender coconut that we are familiar with, is nothing but free nuclear endosperm (made up of thousands of nuclei) and the surrounding white kernel is the cellular endosperm.
  • 35. Embryo Development  Embryo develops at the micropylar end of the embryo sac where the zygote is situated. Most zygotes divide only after certain amount of endosperm is formed. This is an adaptation to provide assured nutrition to the developing embryo. Though the seeds differ greatly, the early stages of embryo development (embryogeny) are similar in both monocotyledons and dicotyledons. the stages of embryogeny in a dicotyledonous embryo.  The zygote gives rise to the proembryo and subsequently to the globular, heart-shaped and mature embryo.  A typical dicotyledonous embryo consists of an embryonal axis and two cotyledons. The portion of embryonal axis above the level of cotyledons is the epicotyl, which terminates with the plumule or stem tip.  The cylindrical portion below the level of cotyledons is hypocotyl that terminates at its lower end in the radicle or root tip. The root tip is covered with a root cap.  Embryos of monocotyledons (Figure 2.14 b) possess only one cotyledon. 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 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 coleoptile.
  • 36. Seed Development  In angiosperms, the seed is the final product of sexual reproduction. It is often described as a fertilised ovule. Seeds are formed inside fruits. A seed typically consists of seed coat(s), cotyledon(s) and an embryo axis.  The cotyledons of the embryo are simple structures generally thick and swollen due to storage of food reserves (as in legumes).  Mature seeds may be non-albuminous or ex-albuminous :  Non- Albuminous: Seeds have no residual endosperm as it is completely consumed during embryo development (e.g., pea, groundnut).  Albuminous seeds: They retain a part of endosperm as it is not completely used up during embryo development (e.g.. Wheat, maize, barley, castor).  Perispermic Seeds : In some seeds such as black pepper and beet, remnants of nucellus are also persistent. This residual, persistent nucellus is the perisperm.  Integuments of ovules harden as tough protective seed coats.  The micropyle remains as a small pore in the seed coat. This facilitates entry of oxygen and water into the seed during germination. As the seed matures, its water content is reduced and seeds become relatively dry (10-15 per cent moisture by mass).  The general metabolic activity of the embryo slows down. The embryo may enter a state of inactivity called dormancy, or if favourable conditions are available (adequate moisture, oxygen and suitable temperature), they germinate.
  • 37. Fruit Formation  As ovules mature into seeds, the ovary develops into a fruit, i.e., the transformation of ovules into seeds and ovary into fruit proceeds simultaneously.  The wall of the ovary develops into the wall of fruit called pericarp.  The fruits may be fleshy as in guava, orange, mango, etc., or may be dry, as in groundnut, and mustard, etc. Many fruits have evolved mechanisms for dispersal of seeds.  In most plants, by the time the fruit develops from the ovary, other floral parts degenerate and fall off.  However, in a few species such as apple, strawberry, cashew, etc., the thalamus also contributes to fruit formation. Such fruits are called false fruits.  Most fruits however develop only from the ovary and are called true fruits.  Although in most of the species, fruits are the results of fertilisation, there are a few species in which fruits develop without fertilisation. Such fruits are called Parthenocarpic fruits. Banana is ones uchexample.  Parthenocarpy can be induced through the application of growth hormones and such fruits are seedless.
  • 38. APOMIXIS & POLYEMBRYONY  Although seeds, in general are the products of fertilisation, a few flowering plants such as some species of Asteraceae and grasses. have evolved a special mechanism, to produce seeds without fertilisation, called apomixis.  Apomixis is a form of asexual reproduction that mimics sexual reproduction.  In some species. The diploid egg cell is formed without reduction division and develops into the embryo without fertilisation.  Some of the nucellar cells surrounding the embryo sac start dividing, protrude into the embryo sac and develop into the embryos. In such species each ovule contains many embryos. Occurrence of more than one embryo in a seed is referred to as polyembryony.