Pollination

8.1 Types of pollination, their advantages and limitations8.2 Methods of pollination, contrivances of
pollination8.3 Fertilization, structure of dicot and monocot embryo
Pollination is the act of transferring pollen grains from the male anther of a flower to the female stigma. The
goal of every living organism, including plants, is to create offspring for the next generation. One of the ways that
plants can produce offspring is by making seeds.
Pollination, transfer of pollen grains from the stamens, the flower parts that produce them, to the ovule-
bearing organs or to the ovules (seed precursors) themselves. In plants such as conifers and cycads, in which the
ovules are exposed, the pollen is simply caught in a drop of fluid secreted by the ovule.
In flowering plants, however, the ovules are contained within a hollow organ called the pistil, and the pollen
is deposited on the pistil’s receptive surface, the stigma. There the pollen germinates and gives rise to a pollen
tube, which grows down through the pistil toward one of the ovules in its base. In an act of
double fertilization, one of the two sperm cells within the pollen tube fuses with the egg cell of the ovule,
making possible the development of an embryo, and the other cell combines with the two subsidiary sexual
nuclei of the ovule, which initiates formation of a reserve food tissue, the endosperm. The growing ovule then
transforms itself into a seed.
As a prerequisite for fertilization, pollination is essential to the production of fruit and seed crops and plays an
important part in programs designed to improve plants by breeding. Furthermore, studies of pollination are
invaluable for understanding the evolution of flowering plants and their distribution in the world today. In flowering
plants, these are (roughly in order of diminishing importance) insects, wind, birds, mammals, and water.
Types: Self-Pollination And Cross-Pollination: An egg cell in an ovule of a flower may be fertilized by a sperm cell derived
from a pollen grain produced by that same flower or by another flower on the same plant, in either of which two cases
fertilization is said to be due to self-pollination (autogamy); or, the sperm may be derived from pollen originating on a
different plant individual, in which case the process is called cross-pollination (heterogamy).

Many species of plants have developed mechanisms that prevent self-pollination. Some—e.g., date palms (Phoenix dactylifera)
and willows (Salix species)—have become dioecious; that is, some plants produce only “male” (staminate) flowers, with the rest
producing only “female” (pistillate or ovule-producing) ones. In species in which staminate and pistillate flowers are found on the
same individual (monoecious plants) and in those with hermaphroditic flowers (flowers possessing both stamens and pistils), a
common way of preventing self-fertilization is to have the pollen shed either before or after the period during which the stigmas
on the same plant are receptive, a situation known as dichogamy. The more usual form of dichogamy, which is found especially in
such insect-pollinated flowers as fireweed (Epilobium angustifolium) and salvias (Salvia species), is protandry, in which the
stamens ripen before the pistils. Protogyny, the situation in which the pistils mature first, occurs in arum lilies and many wind-
pollinated plants, such as grasses—although several grasses are self-pollinated, including common varieties of wheat, barley, and
oats. Avocado has both protogynous and protandrous varieties, and these often are grown together to encourage cross-fertilization.
A structural feature of flowers that discourages selfing is heterostyly, or variation in the length of the style (neck of the pistil). This
occurs in the common primrose (Primula vulgaris) and species of wood sorrel (Oxalis) and flax. In most British primrose
populations, for example, approximately half the individuals have so-called “pin” flowers, which possess short stamens and a long
style, giving the stigma a position at the flower’s mouth, whereas the other half have “thrum” flowers, in which the style is short
and the stamens are long, forming a “thrumhead” at the opening of the flower. Bees can hardly fail to deposit the pollen they
receive from one type of flower onto the stigmas of the other type. The genetic system that regulates flower structure in these
primroses is so constituted that cross-pollination automatically maintains a 50:50 ratio between pins and thrums. In the flowers
of purple loosestrife (Lythrum salicaria), the stamens and styles are of three different lengths to limit self-fertilization.
Chemical: Chemical self-incompatibility is another device for preventing self-fertilization. In this phenomenon,
which depends on chemical substances within the plant, the pollen may fail to grow on a stigma of the same
flower that produced it or, after germination, the pollen tube may not grow normally down the style to
effect fertilization. The process is controlled genetically; it need not be absolute and can change in degree during
the flowering season. Not surprisingly, chemical incompatibility usually is not found in those plants that have
strong structural or temporal barriers against self-pollination. Formation of one such mechanism during evolution
apparently was enough for most plant species. Mechanisms that permit self-pollination: In many instances,
successful self-pollination takes place at the end of a flower’s life-span if cross-pollination has not occurred. Such
self-pollination may be achieved by curving of stamens or style as occurs, for example, in fireweed. It can be an
evolutionary advantage when animal pollinators are temporarily scarce or when the plants in a population are
widely scattered. Under such circumstances, selfing may tide the species over until better circumstances for
outbreeding arrive. For this reason, selfing is common among annual plants; these often must produce an
abundance of seed for the rapid and massive colonization of any bare ground that becomes available. If, in a given
year, an annual plant were to produce no seed at all, survival of the species might be endangered. A persistent habit
of self-pollination apparently has been adopted successfully by some plant species whose natural pollinators have
died out. Continued selfing also is practiced by many food-crop plants. Some of these plants are cleistogamous,
meaning that the flowers fail to open, an extreme way of ensuring self-pollination. A similar process
is apomixis, the development of an ovule into a seed without fertilization. Apomixis is easily demonstrated in
lawn dandelions, which produce seeds even when stamens and styles are cut off just before the flowers open.
Consistent apomixis has the same pros and cons as continued selfing. The offspring show very little genetic
variability, but there is good survival if the species is well adapted to its habitat and if the environment does not
change.
Advantages of Self-pollination : Self- pollination ensures that recessive characters are eliminated. The wastage of the pollen
grain is very less compared to cross-pollination. In the process of self- pollination, the purity of the race is maintained, as there is
no diversity in the genes. In self- pollination, there is no involvement of external factors like wind, water, and other pollinating
agents. Self-pollination ensures that even a smaller quantity of produced pollen grains from plants have a good success rate in
pollination. Disadvantages: The major disadvantage of Self- pollination is there is no mixing up of genes. Due to which: The
vigour and vitality of the race are reduced. The immunity to diseases is reduced in the resultant offsprings.
Types of Cross-Pollination: The process of cross-pollination requires the help of biotic and abiotic agents like animals, birds,
wind, insects, water and other agents as pollinators. Pollination by Wind- Anemophily: There are only a few flowers that
use wind pollination and their features are greenish, small and odourless flowers. As these flowers do not attract the pollinators,
their energy is not used for making colourful petals. This type of pollination usually occurs when plants lack flowers with nectar
and other features including inconspicuous. The male parts of the Anemophilous flowers tend to produce very large quantities of
pollen and the stigma, the female reproductive part of a flower are very large, sticky and feathery to extend completely outside
the flower. Thus the pollen is more likely to reach them. Coconut, palm, maize, grasses and all gymnosperms are the best
examples of wind-pollinated plants. Pollination by Animals – Zoophily: Animals play an important role in plant reproduction.
They help in seed dispersal. Entemophily: By insects. ,Artificial Pollination – Anthropophily: Artificial pollination is done by
human beings. This process is also called as the Anthropophily. If there are any difficulties in the pollination process
through abiotic female flowers. Hybridization techniques are also used in this process. Advantages Cross-pollination: The
produced seeds are good in vigour and vitality. All unisexual plants can reproduce through the process of Cross-pollination. The
recessive characters in the lineage are eliminated as a result of genetic recombination. This process improves the immunity of
the offsprings towards the diseases and other environmental factors. Cross-pollination introduces new genes into a sequence of
species and this is mainly due to the fertilization between genetically different gametes. Disadvantages: In this process, there is
a great wastage of pollen grains. Due to genetic recombination during meiosis, there are chances of eliminations of good
qualities and additions of unwanted characteristics in offspring.
FERTILIZATION: In Gymnosperms, pollen grains usually come in contact with micropyle or nucellus. However, in
angiosperms a special landing place for visiting pollen grains is produced called as 'stigma' When pollen grains are
discharged from anthers they shows considerable resistance to the environmental conditions upto fertilization.
Definition : "The process by which there is fusion of male and female gametes in sexual reproduction, is
known as fertilization." The fertilization results in the formation of an embryo, in most of the plants. However, in
angiosperms double fertilization takes place for the development of endosperm.
Germination of pollen grains: pollen grains stigma produce poisonous substance or may become dry. The time Pollen grains
usually germinate on the stigma of same species. In case of foreign required for germination of pollen grains on stigma varies
from plant to plant. . Saccharum officinarum - within a minute . Beta vulgaris (Beet) - 2 hours Ex.: 1. Sorghum vulgare and . Zea
mays - 5 minutes . Carrya elliptica - 2 days. The germination of separate results in the formation of pollen tube. Pollen tube
is produced from intine and comes out through germpore. Most of the pollen grains are "monosiphonous" (i.e. only one germ
tube is produced during germination). However pollen grains of Malvaceae and Cucurbitaceae are 'polysiphonous'. Stener (1925)
reported about 14 pollen tubes in Malva neglecta. Sometimes same pollen tube shows framification'i.e. branching (Acacia).
Growth of Pollen tube :After the tube has emerged from the pollen grains it goes downward towards the ovary and ovules. The
length of pollen tube depends upon the length of style. It is very short if the stigma is sessile, while it is very long if style is very
long. In Zea mays about 50 cm. pollen tube is recorded and it is called as 'silk. On the basis of presence or absence of
transmiting tissue' Hanf (1935) classified styles into 3 types: 1. Open style :- Where there is wide stylar canal and inner
epidermis helps in the nutrition and conduction of pollen tube. e.g. Family – Papavaraccae 2. Half - closed style :- In this type
canal is surrounded by. Rudimentary transmitting tissue. It is about 2-3 layered and made up of glandular cell. e.g. Cactaceae
3. Closed style :- In this case open channel is absent instead of which a solid core of elongated cells rich in protoplast is present.
e.g. Datura, Gossypium.
Rate of growth of pollen tube : The rate of growth of pollen tube is usually affected by a) Environmental factors, by
Compatibility
a) Environmental factors : Amongst all environmental factors temperature plays vital role. Hofmeistar (1861) had observed that
in Crocus in warm moist air and bright sunshine the pollen tubes can be seen in micropyle within 24 hours. b) Compatibility :
Compatibility between gametetophyte and style sporophytic tissue also affects rate of growth of pollen tube. In compatible species
rate is rapid while in case of incompatible species rate of growth is extremely poor.
Entry of pollen tube and discharge of pollen tube content: Pollen tubes enter inside the ovule by three different methods.
a) Porogamy : When pollen tube enters in ovule directly through micropyle, the condition is known as porogamy. e.g. Most of the angiosperms. b ) Chalazogamy : When
pollen tubes enter inside the ovule through chalazal end then it is called as chalazogamy was first reported by Treub (1891) Swamy (1948) reported Chalazogamy in Casuarina
equisetifolia. c) Mesogamy: When pollen tube enters inside ovule directly through funiculus or integuments then it is known as misogamy. e.g. Cucurbita - Longo (1901)
Finally, pollen tube enters inside the ovule near egg apparatus. The tip of pollen tube bursts and two sperms gets released inside the embryo sac.
Mechanism of fertilization: When male gametes are discharged inside the embryo sac these are having distinct cytoplasmic
sheath. However, in old days these were considered to be only male These two male gametes are found to be unequal in size in
Vinca by Finn (1928). He suggested that, whather smaller gamete is fussing with egg or larger gamete is fussing Cardiospermum
and reported that male gamete fussing with an egg is having a blunt with egg is very difficult ot understand. Cadry (1946) studied
male posterior end and circular or pointed anterior end. The satisfactory explanation regarding the mechanism of fertilization has
been given by, (1) Girassimora (1933) in Crepis capillaries (ii) Whylie's (1928, 1941) in Vallisneria americana. According to
Girassimora certain difficulties arise during the study of process of fertilization. There are as follows :-
1. Along with Sperms some darkly stained bodies are also discharged by pollen tube near egg apparatus. 2. Sometimes synergid
cells degenerate before fertilization and discharge their cytoplasm and nucleus near egg apparatus. 3. The time required for the
process of fertilization is very short and hence it is very difficult to capture that particular movement of fertilization.
Thus very little information is available on the process of fertilization.
According to Girrasimova the mechanism of fertilization is as follows:
When male gametes come in contact with egg nucleus or female gamete, it appears like a thread rolled in to a boll. Later on
this male gamete uncoils and lies on the surface of egg nucleus. Slowly this male gamete immerses in side the egg
completely. Once it comes inside the egg the breaking of chromatin material of male gamete starts. Simultaneously, nucleus
of male gamete begins to appear. Slowly this nucleolus enlarges in size and chromatin material slowly becomes porous.
While chromatin material of egg nucleus becomes distinct.
Finally, the nucleolus of male and female gametes come together and fuse. The fusion of male and female nucleoli ends in
the process of fertilization. The second male gamete combines with secondary nucleolus to form primary
endosperm cell, by the same method. However, time required for this fusion is less than fusion of male gamete
with secondary nucleus is known as double fertilization of triple fusion.
 Double fertilization: Nawashin (1898) was the first man to suggest that, both the sperms released by pollen tube are involved
in fertilization. They fertilize two different elements of the embyo sac. The phenomenon on is unique to angiosperms and is
called as double fertilization
The nucleus of one sperm fuses with the egg nucleus (syngamy) and that of second nucleus fuses with secondary nucleus. the
second fertilization involves fusion of 3 nuclei. The second fertilization involves In most of the plants this secondary nucleus
possesses two polar nuclei. Further, the secondary nucleus may possesses one, three, four or many nuclei. fusion of 3 nuclei. The
phenomenon is called as triple fusion. However in some plants
Triple fusion: After the fusion of one of the male gamete with egg nucleus, one male gamete is still secondary nucleus is diploid
(2n) because it is formed by migration of one nucleus from present there. This remaining male gamete fuses with secondary
nucleus, The migrated from polar region of embryo sac hence called polar nuclei. These two haploid upper side of other form
lower side of embryo sac. Both these nuclei are haploid and polar nuclei after migration at centre of embryo sac, fuses together
and thus becomes diploid. This diploid structure is called as secondary nucleus, One male gamete in haploid (n) and the secondary
nucleus is diploid (2n). The fusion of male gamete and secondary nucleus i.e. n + 2n forms 3n structure hence this fusion is called
as triple fusion. The triple fusion results in the formation of an endosperm, The endosperm is nutritive in function, it provides
nutrition to embryo during early stages of development from zygote to globular embryo.
Significance of double fertilization: Double fertilization process plays significant role in the formation of seeds in the
angiospermic plants. As this process results into production of zygote and primary endosperm nucleus in the fertilized embryo
sac (ovule) fertilized ovules develops into seed. Because Diploid zygote by the process of embryogenesis develops embryo
while primary endosperm nucleus develops a tissue packed with reserve food i.e. endosperm. This endosperm provides
nourishment to the developing embryo as well reserve the food for the maintenance of the embryo during resting or dormancy
period as well as it provides nourishment to germinating embryo. Hence double fertilization has a great significance in inducing
and establishing seed habit in the plants.
ENDOSPERM : Endosperm is the nutritive tissue derived by the fusion of secondary nucleus and second male
gamete. The main function of the endosperm is better nutrition of developing embryo. The endosperm is also present
in Gymnosperms, which differs from the endosperm of angiosperms in its mode of development. In gymnosperms
endosperm is produced before fertilization and is totally maternal while it is produced after fertilization in
angiosperms and hence it is maternal as well as paternal. The development of endosperm after double fertilization
is the distinguishing feature of angiosperms. However, there are few angiosperms which do not form endosperm
are the members of families of Orchidaceae, Podostemonaceoe and Trapacecae. In those angiosperms where
endosperm is developed, it may or may not preserved in metered seeds.
On the basis of presence or absence of endosperm in matured seeds the seeds are broadly classified into two groups,
Endospermic seeds : These seeds are also known as albulminous seeds. In endospermic seeds the developing
embryo never utilizes whole endosperm tissue. Thus in these seeds some amount of endosperm is preserved in
mature seeds. E.g. Castor bean, Seasamum, Poaceae. Non-endospermic seeds : In non endospermic seeds are also
known as exalbuminous seeds. In ex-albuminous seeds the developing embryo utilizes whole endosperm tissue.
Thus in these seeds endosperm is not preserved, it remains in the form of this layer known as 'perisperm’ e.g. All
legumes.
Types of Endosperms: On the basis of mode of development endosperm is classified in to three different types. 1.
Nuclear endosperm 2. Cellular endosperm 3. Helobial endosperm
According to Davis (1966) out of 288 families, about 161 families show nuclear endosperm, 72 families show
cellular endosperm and 17 families possess helobial endosperm
1. Nuclear endosperm : In nuclear endosperm first and other subsequent divisions of primary endosperm cell are
not followed by cell wall formation. The nuclear endosperm is very common in angiosperms and found to occur in
161 families. The development of nuclear endosperm has been well illustrated in Acalypha indica by Johri and Kapil
(1953). According to them the development is as follows. The primary endosperm cell undergoes many free nuclear
divisions. First this cell begins to enlarge in its size. The enlargement is followed by the nuclear divisions of primary
endosperm cell. Slowly central vacuole begins to appear. Thus all the nuclei gets pushed towards periphery or
wallword side. Finally large number of nuclei are produced in a common sheath of cytoplasm and remains at the
peripheral side. According to Johri and Kapil first few nuclear divisions or primary endosperm cell are 'synchronous
(regular) while later on the nuclear divisions by 'asynchronous' (irregular). In case of coconut, the cell wall formation
is incomplete. In this plant all nuclei are utilized for the cell wall formation at the wall word direction producing
coconut meat while central liquid commonly known as coconut milk is without nuclei.
2. Cellular endosperm : In cellular endosperm, the first and other subsequent divisions of primary endosperm cell are followed
by cell wall formation. The cell wall formation since from the beginning makes the embryo sac cellular throughout its
development. Usually the development of cellular endosperm is classified into different sub - types, by Schnarf (1929). This
classification is made on the basis of type of cell wall formation during early stages. We can study with the help of following
examples. a) In Drymis winteri the development of endosperm has been studied by Bhandari and Venkatraman (1968). According
to them in this plant first and other few subsequent divisions of primary endosperm cells are transverse. This results in the
formation of filamentous endosperm. Further divisions of the endosperms cells are irregular. b) In Adoxa the development of
endosperm has been studies by Lagerberg (1909). In this case, the first and second division of primary endosperm is vertical
making four vertical or cylindrical cells. The third division is transverse and results in eight cells and arranged in two tires. Each
tire possesses four cells. The fourth division is also transverse making sixteen celled endosperm tissue. Further divisions are
irregular and that fulfills the whole embryo sac area.
3. Helobial endosperm : This is intermediate type of endosperm between the nuclear and cellular types. This type is
most frequent in the order Helobiales. In this case the first division of primary endosperm cell is followed by
transverse wall producing micropylar and chalazal cell. Subsequently normal free nuclear divisions takes place in the
micropylar cell as well as chalazal cell. But as a rule the main body of endosperm is formed from the micropylar cell
and chalazal cell degenerates or becomes non functional. It is also studied in Asphoedalus tenuifolius. Usually it is
also always in monocots.
An embryo is structure developed from a diploid zygote by repeated mitotic division and through the embryogenesis
capable form emerging into a new plant. The fusion of male gamete with egg nucleus results in the formation of
zygote. This fusion is called as syngamy. The zygote after repeated division develops an embryo. A typical embryo
comprises an embryonal axis, cotyledons, above embryonal axis epicotyls and below hypocotyls. Epicotyl converts
into plumule i.e. embryonic shoot and the hypocotyls at its base has radicle i.e. embryonic root. The embryo of
monocot differs from dicot having only one cotyledons, in dicots cotyledons are two in number. The process of
embryo development is called embryogeny. There are some difference in developmental patterns in embryogeny of
dicots and monocots. In most of the angiosperms zygote divides transversely resulting in small apical cell ca, and large
basal cell cb at micropylar end. Rarely the division of zygote is vertical e.g. Loranthaceae. The variations in the
developmental pattern of embryo during early embryogeny are common in monocots and dicots.
Dicot Embryo:
1. Basal cell forms a 6-10 celled suspensor. 2. Terminal cell produces embryo except the radicle. 3. The first division
of terminal cell is generally longitudinal. 4. It has two cotyledons. 5. plumule is terminal and lies in between the two
elongated cotyledons.

Monocot Embryo: 1. Basal cell produces a single celled suspensor. 2. it forms the whole of the embryo. 3. It is
transverse. 4. There is a single cotyledons. 5. plumule appears lateral due to excessive growth of the single cotyledon.
Dicot Embryo: Atypical Dicot embryo consists of Embryonal axis, epicotyls (Plumule), Hypocotyl (radicle and 2 cotyledons.
Monocot embryo having only one cotyledon.
Seed Formation: Definition: true seed is a fertilised mature ovule, which encloses the embryonic plant, lot of food and protective seed coat,
After fertilization ovules get converted into seed. After fertilization funicle called stalk. After fertilization integuments converted into seed coat (testa), After fertilization nucellus becomes perisperm and After fertilization egg cell develops into an embryo.
Structure of Mature seed: 1. Testa: seeds are enclosed in seed coat known as testa. 2. Embryo: consists 2 large green cotyledons with embryonic axis, having embryonic root (radical) and embryonic shoot (plumule). 3. Hypocotyl: The part of an embryonic axis lying between the radicle and the point of attachment to the axis to the cotyledons called hypocotyl. 4. Epicotyl: The part of axis
between plumule and cotyledon is called epicotyls.. 5. Funicle, The seeds are attached to the fruit by a stalk called funiculus. The scar of it called hilum. 6. The micropyle is a small pore, through which water enters or absorbed during germination. 7. Endosperm: the seed coat encloses white fleshy mass known as endosperm. It is reserve food e. Castor (albuminous, and pea, sunflower,
gram is exalbuminous.