Pollination

Pollination is the process by which pollen is transferred to the female

reproductive organs of a plant, thereby enabling fertilization to take place. Like
all living organisms, seed plants have a single major purpose: to pass their
genetic information on to the next generation. The reproductive unit is the
seed, and pollination is an essential step in the production of seeds in all
spermatophytes (seed plants). For the process of pollination to be successful, a
pollen grain produced by the anther, the male part of a flower, must be
transferred to a stigma, the female part of the flower, of a plant of the same
species. The process is rather different in angiosperms (flowering plants) from
what it is in gymnosperms (other seed plants). In angiosperms, after the pollen
grain has landed on the stigma, it creates a pollen tube which grows down the
style until it reaches the ovary. Sperm cells from the pollen grain then move
along the pollen tube, enter the egg cell through the micropyle and fertilise it,
resulting in the production of a seed.
A successful angiosperm pollen grain (gametophyte) containing the male
gametes is transported to the stigma, where it germinates and its pollen tube
grows down the style to the ovary. Its two gametes travel down the tube to
where the gametophyte(s) containing the female gametes are held within the
carpel. One nucleus fuses with the polar bodies to produce the endosperm
tissues, and the other with the ovule to produce the embryo Hence the term:
"double fertilization"

Introduction
In gymnosperms, the ovule is not contained in a carpel, but exposed on the
surface of a dedicated support organ, such as the scale of a cone, so that the
penetration of carpel tissue is unnecessary. Details of the process vary
according to the division of gymnosperms in question. Two main modes of
fertilization are found in gymnosperms. Cycads and Ginkgo have motile sperm
that swim directly to the egg inside the ovule, whereas conifers and
gnetophytes have sperm that are unable to swim but are conveyed to the egg
along a pollen tube.
The study of pollination brings together many disciplines, such as botany,
horticulture, entomology, and ecology. The pollination process as an
interaction between flower and pollen vector was first addressed in the 18th
century by Christian Konrad Sprengel . It is important in horticulture and
agriculture, because fruiting is dependent on fertilization: the result of
pollination. The study of pollination by insects is known as anthecology.
Process
Pollen germination has three stages; hydration, activation and pollen tube
emergence. The pollen grain is severely dehydrated so that its mass is reduced
enabling it to be more easily transported from flower to flower. Germination
only takes place after rehydration, ensuring that premature germination does
not take place in the anther. Hydration allows the plasma membrane of the
pollen grain to reform into its normal bilayer organization providing an
effective osmotic membrane. Activation involves the development of actin
filaments throughout the cytoplasm of the cell, which eventually become
concentrated at the point from which the pollen tube will emerge. Hydration
and activation continue as the pollen tube begins to grow.
In conifers, the reproductive structures are borne on cones. The cones are
either pollen cones (male) or ovulate cones (female), but some species are
monoecious and others dioecious. A pollen cone contains hundreds of
microsporangia carried on (or borne on) reproductive structures called
sporophylls. Spore mother cells in the microsporangia divide by meiosis to
form haploid microspores that develop further by two mitotic divisions into
immature male gametophytes (pollen grains). The four resulting cells consist of
a large tube cell that forms the pollen tube, a generative cell that will produce
two sperm by mitosis, and two prothallial that degenerate. These cells
comprise a very reduced microgametophyte, that is contained within the
resistant wall of the pollen grain.
The pollen grains are dispersed by the wind to the female, ovulate cone that is
made up of many overlapping scales (sporophylls, and thus megasporophylls),
each protecting two ovules, each of which consists of a megasporangium (the
nucellus) wrapped in two layers of tissue, the integument and the cupule, that
were derived from highly modified branches of ancestral gymnosperms. When
a pollen grain lands close enough to the tip of an ovule, it is drawn in through
the micropyle (a pore in the integuments covering the tip of the ovule) often
by means of a drop of liquid known as a pollination drop.
The pollen enters a pollen chamber close to the nucellus, and there it may wait
for a year before it germinates and forms a pollen tube that grows through the
wall of the megasporangium (=nucellus) where fertilisation takes place. During
this time, the megaspore mother cell divides by meiosis to form four haploid
cells, three of which degenerate. The surviving one develops as a megaspore
and divides repeatedly to form an immature female gametophyte (egg sac).
Two or three archegonia containing an egg then develop inside the
gametophyte. Meanwhile, in the spring of the second year two sperm cells are
produced by mitosis of the body cell of the male gametophyte. The pollen tube
elongates and pierces and grows through the megasporangium wall and
delivers the sperm cells to the female gametophyte inside. Fertilisation takes
place when the nucleus of one of the sperm cells enters the egg cell in the
megagametophyte’sarchegonium.
In flowering plants, the anthers of the flower produce microspores by meiosis.
These undergo mitosis to form male gametophytes, each of which contains
two haploid cells. Meanwhile, the ovules produce megaspores by meiosis,
further division of these form the female gametophytes, which are very
strongly reduced, each consisting only of a few cells, one of which is the egg.
When a pollen grain adheres to the stigma of a carpel it germinates,
developing a pollen tube that grows through the tissues of the style, entering
the ovule through the micropyle. When the tube reaches the egg sac, two
sperm cells pass through it into the female gametophyte and fertilisation takes
place
Fertilization

Types of Pollination
Self-pollination and Cross Pollination (Xenogamy).
Self Pollination is further divided into Autogamy and Geitonogamy. Depending
on agent of Pollination, pollination can be classified into abiotic pollination and
biotic pollination.
• Self Pollination is the type of Pollination in which pollen grains are
transferred from anther to the stigma of the same flower (Autogamy) or pollen
grains are transferred from anther to the stigma of different flower of the
same plant (Geitonogamy).
• Cross Pollination or Xenogamy is the type of pollination in which pollen
grains are transferred from anther to the stigma of a different plant.
On the Basis of Pollinating Agent

• Abiotic pollination refers to situations where pollination is mediated without

the involvement of other organisms. The most common form of abiotic
pollination, anemophily, is pollination by wind. Wind pollination is very
imprecise, with a minute proportion of pollen grains landing by chance on a
suitable receptive stigma, the rest being wasted in the environment. This form
of pollination is used by grasses, most conifers, and many deciduous trees.
Hydrophily is pollination by water, and occurs in aquatic plants which release
their pollen directly into the surrounding water. About 80% of all plant
pollination is biotic. In gymnosperms, biotic pollination is generally incidental
when it occurs, though some gymnosperms and their pollinators are mutually
adapted for pollination. The best-known examples probably are members of
the order Cycadales and associated species of beetles.
Of the abiotically pollinated species of plant, 98% are anemophilous and 2%
hydrophilous, their pollen being transported by water.It is thought that among
angiosperms, entomophily is the primitive state; this is indicated by the
vestigial nectaries in the wind-pollinated Urtica and other plants, and the
presence of fragrances in some of these plants. Of the angiosperms, grasses,
sedges, rushes and catkin-bearing plants are in general wind pollinated. Other
flowering plants are mostly biotic, the pollen being carried by animal vectors.
However a number of plants in multiple families have secondarily adopted
wind pollination in contrast to other members of their groups. Some plants are
intermediate between the two pollination methods. common heather is
regularly pollinated by insects, but produce clouds of pollen and some wind
pollination is inevitable, and the hoary plantain is primarily wind pollinated,
but is also visited by insects which pollinate it.
• Biotic
More commonly, the process of pollination requires pollinators: organisms
that carry or move the pollen grains from the anther of one flower to the
receptive part of the carpel or pistil (stigma) of another. This is biotic
pollination.[9] The various flower traits (and combinations thereof) that
differentially attract one type of pollinator or another are known as pollination
syndromes.[10] At least 100,000 species of animal, and possibly as many as
200,000, act as pollinators of the estimated 250,000 species of flowering plants
in the world.[7] The majority of these pollinators are insects, but about 1,500
species of birds and mammals have been reported to visit flowers and may
transfer pollen between them. Besides birds and bats which are the most
frequent visitors, these include monkeys, lemurs, squirrels, rodents and
possums.Entomophily, pollination by insects, often occurs on plants that have
developed colored petals and a strong scent to attract insects such as, bees,
wasps and occasionally ants (Hymenoptera), beetles (Coleoptera), moths and
butterflies (Lepidoptera), and flies (Diptera)

The existence of insect pollination dates back to the dinosaurera.Inzoophily,

pollination is performed by vertebrates such as birds and bats, particularly,
hummingbirds, sunbirds, spiderhunters, honeyeaters, and fruit bats.
Ornithophily or bird pollination is the pollination of flowering plants by birds.
Chiropterophily or bat pollination is the pollination of flowering plants by bats.
Plants adapted to use bats or moths as pollinators typically have white petals,
strong scent and flower at night, whereas plants that use birds as pollinators
tend to produce copious nectar and have red petals.Insect pollinators such as
honey bees (Apismellifera), bumblebees (Bombusterrestris),and butterflies
(Thymelicusflavus) have been observed to engage in flower constancy, which
means they are more likely to transfer pollen to other conspecific plants. This
can be beneficial for the pollinators, as flower constancy prevents the loss of
pollen during interspecific flights and pollinators from clogging stigmas with
pollen of other flower species. It also improves the probability that the
pollinator will find productive flowers easily accessible and recognisable by
familiar clues.

Mechanism
Pollination can be accomplished by cross-pollination or by self-pollination:
Cross-pollination, also called allogamy, occurs when pollen is delivered from
the stamen of one flower to the stigma of a flower on another plant of the
same species. Plants adapted for cross-pollination have several mechanisms to
prevent self-pollination; the reproductive organs may be arranged in such a
way that self-fertilisation is unlikely, or the stamens and carpels may mature at
different timesModes of Cross Pollination:
The agencies which transfer pollen grains from anthers of one flower to the
stigma of a different flowers are as follows: WIND (Anemophily), WATER
(Hydrophily), INSECTS (Entomophily), BIRDS (Ornithophily)' and BATS
(Cheiropterophily).
(1) Anemophily:
Anemophilous plants produce enormous amount of pollen grains: A single
plant of Mercurialis annually has been estimated to produce 1,352,000,000
pollen grains. Anemophilous plants bear small and inconspicuous flower. The
pollen grains are small, light, smooth and dry. Pollen of some plants are said to
be blown to 1,300 km. In some plants as Pinus, pollen grains are winged.
The flowers are usually unisexual in some plants e.g. Mulberry is borne in
independent catkins which can sway freely and shake off their pollen in air.
The flowers may be borne on long axis (as in grasses) much above the leaves.
The anther is versatile so as to oscillate in all directions at the tip of filament. In
Urticaceae filaments are very long. Anempohilous flowers have adequate
devices to catch the air-borne-pollen grains with utmost efficiency. For this the
stigma is usually large and feathery (as in grasses) and brush like as in Typha.

(2) Hydrophily:
It is of two types:
(a) Hypohydrogamy:
Includes plants which are pollinated inside the water, e.g. Ceratophyllum,
Najas.
(b) Epihydrogamy:
Vallisneriaspiralis (ribbon weed) is a submerged dioecious plant. The flowers
are borne under water. When mature, the male flower get detached from the
parent plant and float on the surface of water. The pistillate flowers also
develop under water, at the time of pollination, they are brought to the
surface by their long and slender stalks. As it arrives on the surface it forms a
cuplike depression. If male flowers floating on water get lodged into the
depression, the pollination takes place. After pollination, the stalk of the
pistillate flower undergoes spiral torsion bringing the pollinated flower under
water once more.

(3) Entomophily:
Some of the insects which help in pollination are bees, flies, wasps, moths and
beetles. Bees, flies and beetles visit flowers which open after sunset. Bees
probably carry out 80% of all pollination done by insects. Bee pollinated
flowers are coloured, possess special smell and/or produce nectar. Pollen
grains are sticky or with spinousexine. Also the stigma is sticky and bees are
colour blind for red.

(4) Ornithophily:
Tiny birds like humming birds and honey thrushes (hardly 1 inch long) feeds on
the nectar of flower like Bignonia, Erythrina is visited by crows.

(5) Chiropteriphily:
Bauhinia megalandra of Java and Anthocephalus are pollinated by bats.

(6) Malcophily:
Many aroids which are usually pollinated by Diptera are also pollinated by snails.
• Self-pollination occurs when pollen from one flower pollinates the same flower or other
flowers of the same individual. It is thought to have evolved under conditions when
pollinators were not reliable vectors for pollen transport, and is most often seen in short-
lived annual species and plants that colonize new locations. Self-pollination may include
autogamy, where pollen is transferred to the female part of the same flower; or
geitonogamy, when pollen is transferred to another flower on the same plant. Plants
adapted to self-fertilize often have similar stamen and carpel lengths. Plants that can
pollinate themselves and produce viable offspring are called self-fertile. Plants that cannot
fertilize themselves are called self-sterile, a condition which mandates cross-pollination for
the production of offspring.
• Cleistogamy: is self-pollination that occurs before the flower opens. The pollen is released
from the anther within the flower or the pollen on the anther grows a tube down the style
to the ovules. It is a type of sexual breeding, in contrast to asexual systems such as apomixis.
Some cleistogamous flowers never open, in contrast to chasmogamous flowers that open
and are then pollinated. Cleistogamous flowers are by necessity found on self-compatible or
self-fertile plants.[23] Although certain orchids and grasses are entirely cleistogamous, other
plants resort to this strategy under adverse conditions. Often there may be a mixture of
both cleistogamous and chasmogamous flowers, sometimes on different parts of the plant
and sometimes in mixed inflorescences. The ground bean produces cleistogamous flowers
below ground, and mixed cleistogamous and chasmogamous flowers above.
Pollen vectors
Biotic pollen vectors are animals, usually insects, but also reptiles, birds,
mammals, and sundry others, that routinely transport pollen and play a role in
pollination. This is usually as a result of their activities when visiting plants for
feeding, breeding or shelter. The pollen adheres to the vector's body parts
such as face, legs, mouthparts, hair, feathers, and moist spots; depending on
the particular vector. Such transport is vital to the pollination of many plant
species.
Any kind of animal that often visits or encounters flowers is likely to be a
pollen vector to some extent. For example, a crab spider that stops at one
flower for a time and then moves on, might carry pollen incidentally, but most
pollen vectors of significant interest are those that routinely visit the flowers
for some functional activity. They might feed on pollen, or plant organs, or on
plant secretions such as nectar, and carry out acts of pollination on the way.
Many plants bear flowers that favour certain types of pollinator over all others.
This need not always be an effective strategy, because some flowers that are
of such a shape that they favor pollinators that pass by their anthers and
stigmata on the way to the nectar, may get robbed by ants that are small
enough to bypass the normal channels, or by short-tongued bees that bite
through the bases of deep corolla tubes to extract nectar at the end opposite
to the anthers and stigma.
Some pollinator species can show huge variation in pollination effectiveness
because their ability to carry pollen is impacted by some morphological trait.
This is the case in the white-lined sphinx moth, in which short-tongued morphs
collect pollen on their heads but long-tongued morphs do not carry any pollen.
Some flowers have specialized mechanisms to trap pollinators to increase
effectiveness. Other flowers will attract pollinators by odor. For example, bee
species such as Euglossacordata are attracted to orchids this way, and it has
been suggested that the bees will become intoxicated during these visits to the
orchid flowers, which last up to 90 minutes. However, in general, plants that
rely on pollen vectors tend to be adapted to their particular type of vector, for
example day-pollinated species tend to be brightly coloured, but if they are
pollinated largely by birds or specialist mammals, they tend to be larger and
have larger nectar rewards than species that are strictly insect-pollinated. They
also tend to spread their rewards over longer periods, having long flowering
seasons; their specialist pollinators would be likely to starve if the pollination
season were too short.

CONCLUSIONS
Pollination management is a branch of agriculture that seeks to protect and
enhance present pollinators and often involves the culture and addition of
pollinators in monoculture situations, such as commercial fruit orchards. The
largest managed pollination event in the world is in Californianalmond
orchards, where nearly half (about one million hives) of the US honey bees are
trucked to the almond orchards each spring. New York's apple crop requires
about 30,000 hives; Maine's blueberry crop uses about 50,000 hives each year.
Bees are also brought to commercial plantings of cucumbers, squash, melons,
strawberries, and many other crops. Honey bees are not the only managed
pollinators: a few other species of bees are also raised as pollinators. The
alfalfa leafcutter bee is an important pollinator for alfalfaseed in western
United States and Canada. Bumblebees are increasingly raised and used
extensively for greenhouse tomatoes and other crops.
The ecological and financial importance of natural pollination by insects to
agricultural crops, improving their quality and quantity, becomes more and
more appreciated and has given rise to new financial opportunities. The
vicinity of a forest or wild grasslands with native pollinators near agricultural
crops, such as apples, almonds or coffee can improve their yield by about 20%.
The benefits of native pollinators may result in forest owners demanding
payment for their contribution in the improved crop results – a simple example
of the economic value of ecological services.