Pollination
Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds. Pollinating agents can be animals such as insects, for example bees, beetles or butterflies; birds, and bats; water; wind; and even plants themselves. Pollinating animals travel from plant to plant carrying pollen on their bodies in a vital interaction that allows the transfer of genetic material critical to the reproductive system of most flowering plants. Self-pollination occurs within a closed flower. Pollination often occurs within a species. When pollination occurs between species, it can produce hybrid offspring in nature and in plant breeding work.
In angiosperms, after the pollen grain has landed on the stigma, it germinates and develops a pollen tube which grows down the style until it reaches an ovary. Its two gametes travel down the tube to where the gametophyte containing the female gametes are held within the carpel. After entering an ovule through the micropyle, one male nucleus fuses with the polar bodies to produce the endosperm tissues, while the other fuses with the egg cell to produce the embryo. Hence the term: "double fertilisation". This process would result in the production of a seed, made of both nutritious tissues and embryo.
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 fertilisation 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.
Pollination research covers various fields, including 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 fertilisation: the result of pollination. The study of pollination by insects is known as anthecology. There are also studies in economics that look at the positives and negatives of pollination, focused on bees, and how the process affects the pollinators themselves.
Process of pollination
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 or ovulate cones, but some species are monoecious and others dioecious. A pollen cone contains hundreds of microsporangia carried 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. 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 cells that degenerate. These cells comprise a very reduced microgametophyte, that is contained within the resistant.
The pollen grains are dispersed by the wind to the female, ovulate cone that is made up of many overlapping scales, each protecting two ovules, each of which consists of a megasporangium 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 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 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. 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's archegonium.
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.
Methods
Pollination may be biotic or abiotic. Biotic pollination relies on living pollinators to move the pollen from one flower to another. Abiotic pollination relies on wind, water or even rain. Adding natural habitat areas into farm systems generally improves pollination, as farms that are closer to natural habitat have higher crop yield because they are visited by more pollinators.Biotic pollination
About 80% of angiosperms rely on biotic pollination. : organisms that carry or move the pollen grains from the anther of one flower to the receptive part of the carpel or pistil of another. Between 100,000 and 200,000 species of animal act as pollinators of the world's 250,000 species of flowering plant. The majority of these pollinators are insects, but about 1,500 species of birds and mammals 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, beetles, moths and butterflies, and flies. The existence of insect pollination dates back to the dinosaur era.
Insect pollinators such as honey bees,
bumblebees, and butterflies 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. The primary insect pollinators are hymenopterans, mostly bees, but also including sawflies, ants, and many species of wasps.
Many flowers attract pollinators by odor. For example, orchid bee species such as Euglossa cordata are attracted to orchids this way, and it has been suggested that some orchid species intoxicate bees during visits which can 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 and have little odor, 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. Night-blooming flowers have little color, but are often very aromatic. Plants with vertebrate pollinators 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.
Some flowers have specialized mechanisms to trap pollinators to increase effectiveness, attach pollen to specific body parts, or require specialized behaviors or morphology in order to extract pollen or nectar. One such syndrome is "buzz pollination", where a bee must vibrate at a certain frequency in order to cause pollen to be released from the anthers.
In zoophily, 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.
Mammals are not generally thought of as pollinators, but some rodents, bats and marsupials are significant pollinators and some even specialise in such activities. In South Africa certain species of Protea are adapted to pollination by rodents and elephant shrews. The flowers are borne near the ground, are yeasty smelling, not colourful, and sunbirds reject the nectar with its high xylose content. The mice apparently can digest the xylose and they eat large quantities of the pollen. In Australia pollination by flying, gliding and earthbound mammals has been demonstrated.
Reptile pollinators are known, but they form a minority in most ecological situations. They are most frequent and most ecologically significant in island systems, where insect and sometimes also bird populations may be unstable and less species-rich. Adaptation to a lack of animal food and of predation pressure, might therefore favour reptiles becoming more herbivorous and more inclined to feed on pollen and nectar. Most species of lizards in the families that seem to be significant in pollination seem to carry pollen only incidentally, especially the larger species such as Varanidae and Iguanidae, but especially several species of the Gekkonidae are active pollinators, and so is at least one species of the Lacertidae, Podarcis lilfordi, which pollinates various species, but in particular is the major pollinator of Euphorbia dendroides on various Mediterranean islands.