Carpenter bee with pollen collected from Night-blooming cereus

  Tip of a tulip stamen. Note the pollen grains

Pollination is the process by which pollen is transferred in the reproduction of plants, thereby enabling fertilization and sexual reproduction.

In spite of a common perception that pollen grains are gametes, like the sperm cells of animals, this is incorrect; pollination is a phase in the alternation of generations: each pollen grain is a male haploid plant, a gametophyte, adapted to being transported to the female gametophyte, where it can achieve fertilization by producing the male gamete (or gametes, in the process of double fertilization).

As such the Angiosperm successful pollen grain (gametophyte) containing the male gametes (sperm) gets 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 ovum to produce the embryo^[1][2] Hence the term: "double fertilization".

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.

The receptive part of the carpel is called a stigma in the flowers of angiosperms. The receptive part of the gymnosperm ovule is called the micropyle. Pollination is a necessary step in the reproduction of flowering plants, resulting in the production of offspring that are genetically diverse.

The study of pollination brings together many disciplines, such as botany, horticulture, entomology, and ecology. The pollination process as an interaction between flower and 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, which is the result of pollination.

  Types

  Abiotic

  Abiotic pollination by wind, depicted in Praesidium Sponsaliorum Plantarum by Carl Linnaeus, 1729.

Abiotic pollination refers to situations where pollination is mediated without the involvement of other organisms. Only 10% of flowering plants are pollinated without animal assistance.^[3] The most common form of abiotic pollination, anemophily, is pollination by wind. This form of pollination is predominant in 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. Most conifera are anemophilous; they depend on wind pollination. Of the 20% of abiotically pollinated species, 98% are anemophilous and 2% hydrophilous, being pollinated by water.^{[citation needed]}

  Biotic

  A hummingbird feeding

More commonly, the process of pollination requires pollinators: organisms that carry or move the pollen grains from the anther to the receptive part of the carpel or pistil. This is biotic pollination. The various flower traits (and combinations thereof) that differentially attract one type of pollinator or another are known as pollination syndromes. Roughly 200,000 varieties of animal pollinators are in the wild, most of which are insects.^[3] 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 dinosaur era.^[4] In zoophily, pollination is performed by vertebrates such as birds and bats, particularly, hummingbirds, sunbirds, spiderhunters, honeyeaters, and fruit bats. Plants adapted to using bats or moths as pollinators typically have white petals and a strong scent, while plants that use birds as pollinators tend to develop red petals and rarely develop a scent (few birds rely on a sense of smell to find plant-based food).

Insect pollinators such as honeybees (Apis mellifera),^[5] bumblebees (Bombus terrestris),^[6][7] and butterflies (Thymelicus flavus) ^[8] have been observed to engage in flower constancy, which means they are more likely to transfer pollen to other conspecific plants.^[9] This can be beneficial for the pollenisers, as flower constancy prevents the loss of pollen during interspecific flights and pollinators from clogging stigmas with pollen of other flower species.^[10]

  Mechanics

Pollination can be accomplished by cross-pollination or by self-pollination :

• Cross-pollination, also called allogamy, occurs when pollen is delivered to a flower from a different plant. Plants adapted to outcross or cross-pollinate often have taller stamens than carpels or use other mechanisms to better ensure the spread of pollen to other plants' flowers.

  A European honey bee collects nectar, while pollen collects on its body.

  California High Desert Honey Bees Immersed in Yellow Beavertail Cactus Flower Pollen

• Self-pollination occurs when pollen from one flower pollinates the same flower or other flowers of the same individual.^[11] 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.^[12] Self pollination may include autogamy, where pollen moves 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 by necessity are self-compatible or self-fertile plants.^[13] Many plants are self-incompatible, and these two conditions are end points on a continuum.

  Geranium incanum, like most geraniums and pelargoniums, sheds its anthers, sometimes its stamens as well, as a barrier to self-pollination. This young flower is about to open its anthers, but has not yet fully developed its pistil.

  These Geranium incanum flowers have opened their anthers, but not yet their stigmas. Note the change of colour that signals to pollinators that it is ready for visits.

  This Geranium incanum flower has shed its stamens, and deployed the tips of its pistil without accepting pollen from its own anthers. (It might of course still receive pollen from younger flowers on the same plant.)

Pollination also requires consideration of pollenizers. The terms "pollinator" and "pollenizer" are often confused: a pollinator is the agent that moves the pollen, whether it be bees, flies, bats, moths, or birds; a pollenizer is the plant that serves as the pollen source for other plants. Some plants are self-fertile or self-compatible and can pollinate themselves (e.g., they act as their own pollenizer). Other plants have chemical or physical barriers to self-pollination.

In agriculture and horticulture pollination management, a good pollenizer is a plant that provides compatible, viable and plentiful pollen and blooms at the same time as the plant that is to be pollinated or has pollen that can be stored and used when needed to pollinate the desired flowers. Hybridization is effective pollination between flowers of different species, or between different breeding lines or populations. see also Heterosis.

Peaches are considered self-fertile because a commercial crop can be produced without cross-pollination, though cross-pollination usually gives a better crop. Apples are considered self-incompatible, because a commercial crop must be cross-pollinated. Many commercial fruit tree varieties are grafted clones, genetically identical. An orchard block of apples of one variety is genetically a single plant. Many growers now consider this a mistake. One means of correcting this mistake is to graft a limb of an appropriate pollenizer (generally a variety of crabapple) every six trees or so.^{[citation needed]}

  The wasp Mischocyttarus rotundicollis transporting pollen grains of Schinus terebinthifolius

  Pollen vectors

Pollen vectors are animals, usually insects, that transport pollen of plants when using the flowers for feeding, breeding or hiding. The pollen is found adhered to insect's body parts such as face, legs and mouthparts or to mammal's hairs, resulting or helping in the pollination of many plant species. An example are wasps, which can transport pollen and contribute for the pollination of several plant species, being potential or even efficient pollinators.^[14]

  Evolution of plant/pollinator interactions

The first fossil record for abiotic pollination is from fern-like plants in the late Carboniferous period. Gymnosperms show evidence for biotic pollination as early as the Triassic period. Many fossilized pollen grains show characteristics similar to the biotically dispersed pollen today. Furthermore, the gut contents, wing structures, and mouthpart morphologies of fossilized beetles and flies suggest that they acted as early pollinators. The association between beetles and angiosperms during the early Cretaceous period led to parallel radiations of angiosperms and insects into the late Cretaceous. The evolution of nectaries in late Cretaceous flowers signals the beginning of the mutualism between hymenopterans and angiosperms.

  In agriculture

  An Andrena bee collects pollen among the stamens of a rose. The female carpel structure appears rough and globular to the left. The bee's stash of pollen is on its hind leg.

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 Californian almond 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 alfalfa seed in western United States and Canada. Bumblebees are increasingly raised and used extensively for greenhouse tomatoes and other crops.

  Well-pollinated blackberry blossom begins to develop fruit. Each incipient drupelet has its own stigma and good pollination requires the delivery of many grains of pollen to the flower so that all drupelets develop.

  Blueberries being pollinated by bumblebees. Bumblebee hives need to be bought each year as the queens must hibernate (unlike honey bees). They are used nonetheless as they offer advantages with certain fruits as blueberries (such as the fact that they are active even at colder outdoor ambient temperature).

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.

The American Institute of Biological Sciences reports that native insect pollination saves the United States agricultural economy nearly an estimated $3.1 billion annually through natural crop production;^[15] pollination produces some $40 billion worth of products annually in the United States alone.^[3]

Pollination of food crops has become an environmental issue, due to two trends. The trend to monoculture means that greater concentrations of pollinators are needed at bloom time than ever before, yet the area is forage poor or even deadly to bees for the rest of the season. The other trend is the decline of pollinator populations, due to pesticide misuse and overuse, new diseases and parasites of bees, clearcut logging, decline of beekeeping, suburban development, removal of hedges and other habitat from farms, and public concern about bees. Widespread aerial spraying for mosquitoes due to West Nile fears is causing an acceleration of the loss of pollinators.

The US solution to the pollinator shortage, so far, has been for commercial beekeepers to become pollination contractors and to migrate. Just as the combine harvesters follow the wheat harvest from Texas to Manitoba, beekeepers follow the bloom from south to north, to provide pollination for many different crops.

  Environmental impacts

Loss of pollinators, also known as Pollinator decline (of which colony collapse disorder is perhaps the most well known) has been noticed in recent years.^[16] Observed losses would have significant economic impacts. Possible explanations for pollinator decline include habitat destruction, pesticide, parasitism/diseases, and others.

  See also

Wikimedia Commons has media related to: pollination

• Cheating (biology)
• Fruit tree pollination
• Hand pollination
• Self-pollination
• Domestication
• Pollination syndrome
• Canadian Pollination Initiative
• Hermann Müller (botanist)
• Paul Knuth (botanist)

  References

• Crepet WL, Friis EM, Nixon KC. 1991. Fossil evidence for the evolution of biotic pollination [and discussion]. Philosophical Transactions: Biological Sciences 333(1267) 187-195.
• Dafni, Amots; Kevan, Peter G.; and Husband, Brian C. (2005). Practical Pollination Biology. Enviroquest, Ltd. ISBN 978-0-9680123-0-7.
• Labandeira CC, Kvacek J, & Mostovski MB. 2007. Pollination drops, pollen and insect pollination of Mesozoic gymnosperms. Taxon 56(3) 663-695.
• Sihag, R.C. 1997.Pollination Biology: Basic and Applied Principles. Rajendra Scientific Publishers,Hisar, 210p.

  External links

Look up pollination in Wiktionary, the free dictionary.

• New "Pollinator Garden Wheel" from the National Academies provides information on pollination and tips on building a pollinator-friendly garden.
• Insect Pollination Of Cultivated Crop Plants by S. E. McGregor USDA 1972 (needs updating but still valuable)
• The Pollination Home page
• Pollination in Hydroponics
• Pollination syndromes images at bioimages.vanderbilt.edu

Botany Subdisciplines of botany Ethnobotany Paleobotany Plant anatomy Plant ecology Plant evo-devo Plant morphology Plant physiology Plants Evolutionary history of plants Algae Bryophyte Pteridophyte Gymnosperm Angiosperm Plant parts Flower Fruit Leaf Meristem Root Stem Stoma Vascular tissue Wood Plant cells Cell wall Chlorophyll Chloroplast Photosynthesis Plant hormone Plastid Transpiration Plant reproduction Alternation of generations Gametophyte Plant sexuality Pollen Pollination Seed Spore Sporophyte Plant taxonomy Botanical name Botanical nomenclature Herbarium IAPT ICN Species Plantarum Glossaries Glossary of botanical terms Glossary of plant morphology terms Category Portal