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[[Image:Misc pollen.jpg|thumb|right|250px|SEM image of pollen grains from a variety of common plants: sunflower (Helianthus annuus), morning glory (Ipomoea purpurea), prairie hollyhock (Sidalcea malviflora), oriental lily(Lilium auratum), evening primrose (Oenothera fruticosa), and castor bean (Ricinus communis).]]
Closeup image of a cactus flower and its stamens
Tip of a tulip stamen.
Note the grains of pollen
Pollen sticking to a bee.
Insects involuntarily transporting pollen from flower to flower play an important role in many plants' reproductive cycles.
Marmalade fly sitting on a grey-haired rockrose, its face and legs covered in pollen.
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes (sperm cells). Pollen grains have a hard coat that protects the sperm cells during the process of their movement between the stamens to the pistil of flowering plants or from the male cone to the female cone of coniferous plants. When pollen lands on a compatible pistil of flowering plants, it germinates and produces a pollen tube that transfers the sperm to the ovule of a receptive ovary. The Individual pollen grains are small enough to require magnification to see detail.

The structure of pollen

Pollen itself is not the male gamete, but each pollen grain contains vegetative (non-reproductive) cells (only a single cell in most flowering plants but several in other seed plants) and a generative (reproductive) cell containing two nuclei: a tube nucleus (that produces the pollen tube) and a generative nucleus (that divides to form the two sperm cells). The group of cells is surrounded by a cellulose-rich cell wall called the intine, and a resistant outer wall composed largely of sporopollenin called the exine.

Pollen is produced in the microsporangium (contained in the anther of an angiosperm flower, male cone of a coniferous plant, or male cone of other seed plants). Pollen grains come in a wide variety of shapes (most often spherical), sizes, and surface markings characteristic of the species (see Electron micrograph at top right). Pollen grains of pines, firs, and spruces are winged. The smallest pollen grain, that of the Forget-me-not (Myosotis spp.), is around 6 µm (0.006  mm) in diameter. Wind-borne pollen grains can be as large as about 90-100 µm. The study of pollen is called palynology and is highly useful in paleoecology, paleontology, archeology, and forensics.

In angiosperms, during flower development the anther is composed of a mass of cells that appear undifferentiated, except for a partially differentiated dermis. As the flower develops, four groups of sporogenous cells form with in the anther, the fertile sporogenous cells are surrounded by layers of sterile cells that grow into the wall of the pollen sac, some of the cells grow into nutritive cells that supply nutrition for the microspores that form by meiotic division from the sporogenous cells. Four haploid microspores are produced from each diploid sporogenous cell called a microsporocyte, after meiotic division. After the formation of the four microspores, which are contained by callose walls, the development of the pollen grain walls begins. The callose wall is broken down by an enzyme called callase and the freed pollen grains grow in size and develop their characteristic shape and form a resistant outer wall called the exine and an inner wall called the intine. The exine is what is preserved in the fossil record.

The pollen wall protects the sperm nucleus while the pollen grain is moving from the anther to the stigma, it protects the vital genetic material from drying out and solar radiation. The pollen grain surface is covered with waxes and proteins, which are held in place by structures called sculpture elements on the surface of the grain. The outer pollen wall prevents the pollen grain from shrinking and crushing the genetic material during desiccation and it is composed of two layers. These two layers are the tectum and the foot layer, which is just above the intine. The tectum and foot layer are separated by a region called the columella, which is composed of strengthening rods. The outer wall is constructed with a resistant biopolymer called sporopollenin. The pollen tube passes through the wall by way of structures called apertures.

Pollen aperture are any modification of the wall of the pollen grain. These modifications include thinning, ridges and pores, they serve as an exit for the pollen contents and allow shrinking and swelling of the grain caused by changes in moisture content. Furrows in the pollen grain are called colpi, which along with pores, are a chief criteria for the identifying pollen classes.

Pollen grains may have furrows, the orientation of which (relative to the original tetrad of microspores) classify the pollen as colpate or sulcate. The number of furrows or pores helps classify the flowering plants, with eudicots having three colpi (tricolpate), and other groups having one sulcus.

Except in the case of some submerged aquatic plants, the mature pollen-grain has a double wall, a thin delicate wall of unaltered cellulose (the endospore or intine) and a tough outer cuticularized exospore or exine. The exine often bears spines or warts, or is variously sculptured, and the character of the markings is often of value for identifying genus, species, or even cultivar or individual. In some flowering plants, germination of the pollen grain often begins before it leaves the microsporangium, with the generative cell forming the two sperm cells.

Pollination

The transfer of pollen grains to the female reproductive structure (pistil in angiosperms) is called pollination. This transfer can be mediated by the wind, in which case the plant is described as anemophilous (literally wind-loving). Anemophilous plants typically produce great quantities of very lightweight pollen grains, sometimes with air-sacs. Non-flowering seed plants (e.g. pine trees) are characteristically anemophilous. Anemophilous flowering plants generally have inconspicuous flowers. Entomophilous (literally insect-loving) plants produce pollen that is relatively heavy, sticky and protein-rich, for dispersal by insect pollinators attracted to their flowers. Many insects and some mites are specialized to feed on pollen, and are called palynivores.

In non-flowering seed plants, pollen germinates in the pollen chamber, located beneath and inside the micropyle. A pollen tube is produced, which grows into the nucellus to provide nutrients for the developing sperm cells. Sperm cells of Pinophyta and Gnetophyta are without flagella, and are carried by the pollen tube, while those of Cycadophyta and Ginkgophyta have many flagella.

When placed on the stigma of a flowering plant, under favorable circumstances, a pollen grain puts forth a pollen tube which grows down the tissue of the style to the ovary, and makes its way along the placenta, guided by projections or hairs, to the micropyle of an ovule. The nucleus of the tube cell has meanwhile passed into the tube, as does also the generative nucleus which divides (if it hasn't already) to form two sperm cells. The sperm cells are carried to their destination in the tip of the pollen-tube.

Pollen as a carrier of ecological information in plants

A Russian theoretical biologist, Vigen Geodakyan (Geodakian), has suggested that the quantity of pollen reaching a pistillate flower can transmit ecological information and also regulate evolutionary plasticity in cross-pollinating plants. Plentiful pollen indicates optimum environmental conditions (for example a plant that is situated at the center of its natural range, in ideal growing conditions, with a large number of male plants nearby, and favorable weather conditions), whereas a small amount of pollen indicates extreme conditions (at the borders of its range, with a deficiency of male plants, and adverse weather conditions). Geodakian believes that the quantity of pollen reaching a pistillate flower defines the sex ratio, dispersion and sexual dimorphism of a plant population. High pollen quantity leads to a reduction of these characteristics and stabilization of a population. Small quantity leads to their increase and destabilization of a population.

Dependence of the secondary sex ratio on the amount of fertilizing pollen was confirmed on four dioecious plant species from three families — Rumex acetosa (Polygonaceae), Melandrium album (Cariophyllaceae), Cannabis sativa and Humulus japonicus (Cannabinaceae). (see summary of all these data in review article).

Dependence of offspring phenotype variety on amount of pollen was observed by Ter-Avanesyan in 1949. All three studied species of plants (cotton plant, black-eyed pea, and wheat) showed dependence in the direction forecast by the theory — fertilization with a small amount of pollen resulted in an increase in the diversity of the offspring. Ter-Avanesian writes that as a result of a limited pollination “instead of homogenous sorts we get populations”.

Pollen in the fossil record

Pollen's sporopollenin outer sheath affords it some resistance to the rigours of the fossilisation process that destroy weaker objects; it is also produced in huge quantities. As such, there is an extensive fossil record of pollen grains, often disassociated from their parent plant. The discipline of palynology is devoted to the study of pollen, which can be used both for biostratigraphy and to gain information about the abundance and variety of plants alive — which can itself yield important information about paleoclimates.Pollen is first found in the fossil record in the late Devonian period and increases in abundance until the present day.

Hay fever

Allergy to pollen is called hay fever. Generally pollens that cause allergies are those of anemophilous plants (pollen is dispersed by air currents.) Such plants produce large quantities of lightweight pollen (because wind dispersal is random and the likelihood of one pollen grain landing on another flower is small) which can be carried for great distances and are easily inhaled, bringing it into contact with the sensitive nasal passages.

In the US, people often mistakenly blame the conspicuous goldenrod flower for allergies. Since this plant is entomophilous (its pollen is dispersed by animals), its heavy, sticky pollen does not become independently airborne. Most late summer and fall pollen allergies are probably caused by ragweed, a widespread anemophilous plant.

Arizonamarker was once regarded as a haven for people with pollen allergies, although several ragweed species grow in the desert. However, as suburbs grew and people began establishing irrigated lawns and gardens, more irritating species of ragweed gained a foothold and Arizona lost its claim of freedom from hay fever.

Anemophilous spring blooming plants such as oak, birch, hickory, pecan, and early summer grasses may also induce pollen allergies. Most cultivated plants with showy flowers are entomophilous and do not cause pollen allergies. In the US, oak pollen starts to cause problems for sufferers in February and is gone by the end of April.

Pollen in human diets

A variety of producers have started selling pollen for human consumption, often marketed as a food (rather than a dietary supplement). The largest constituent is carbohydrates, with protein content ranging from 7 to 35 percent depending on the plant species collected by bees.

The U.S. Food and Drug Administration (FDA) has not found any harmful effects of pollen consumption, except from the usual allergies. However, FDA does not allow pollen marketers in the United States to make health claims about their produce, as no scientific basis for these has ever been proved. Furthermore, there are possible dangers not only from allergic reactions but also from contaminants such as pesticides and from fungi and bacteria growth related to poor storage procedures. A manufacturers's claim that pollen collecting helps the bee colonies is also controversial.

Forensic palynology

In forensic biology, pollen can tell a lot about where a person or object has been, because regions of the world, or even more particular locations such a certain set of bushes, will have a distinctive collection of pollen species. Pollen evidence can also reveal the season in which a particular object picked up the pollen. Pollen has been used to trace activity at mass graves in Bosniamarker, catch a burglar who brushed against a Hypericum bush during a crime, and has even been proposed as an additive for bullets to enable tracking them.

See also



References

  1. http://www.geo.arizona.edu/palynology/polkey.html#key
  2. http://www.geo.arizona.edu/palynology/ppapertr.html
  3. ( full text)
  4. Geodakyan V. A. (1977). The Amount of Pollen as a Regulator of Evolutionary Plasticity of Cross-Pollinating Plants. “Doklady Biological Sciences” 234 N 1-6, 193–196.
  5. Соrrеns С. (1922) Geschlechtsbestimmung und Zahlenverhaltnis der Geschlechter beim Sauerampfer (Rumex acetosa). “Biol. Zbl.” 42, 465-480.
  6. Rychlewski J., Kazlmierez Z. (1975) Sex ratio in seeds of Rumex acetosa L. as a result of sparse or abundant pollination. “Acta Biol. Cracov” Scr. Bot., 18, 101-114.
  7. Correns C. (1928) Bestimmung, Vererbung und Verteilung des Geschlechter bei den hoheren Pflanzen. Handb. Vererbungswiss., 2, 1-138.
  8. Mulcahy D. L. (1967) Optimal sex ratio in Silene alba. “Heredity” 22 № 3, 41.
  9. Riede W. (1925) Beitrage zum Geschlechts- und Anpassungs-problem. “Flora” 18/19
  10. Kihara H., Hirayoshi J. (1932) Die Geschlechtschromosomen von Humulus japonicus. Sieb. et. Zuce. In: 8th Congr. Jap. Ass. Adv. Sci., p. 363—367 (cit.: Plant Breeding Abstr., 1934, 5, № 3, p. 248, ref. № 768).
  11. Geodakyan, V.A. & Geodakyan, S.V., (1985) Is there a negative feedback in sex determination? “Zurnal obschej biol.” 46 201-216 (in Russian). ).
  12. Ter-Avanesyan D. V. (1949). Tr. Prikl. Bot, Genet, Selekt., 28 119.
  13. Ter-Avanesian D. V. (1978) Significance of pollen amount for fertilization. “Bull. Torrey Bot. Club.” 105 N 1, 2–8.
  14. Sanford, citing P. Witherell, "Other Products of the Hive," Chapter XVIII, The Hive and the Honey Bee, Dadant & Sons, Inc., Hamilton, IL, 1975
  15. . Document ENY118. Original publication date November 1, 1994. Revised February 1, 1995. Reviewed May 1, 2003.
  16. Newscripts, Chemical & Engineering News, 86, 33, 18 Aug. 2008, p. 88


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Clumps of yellow pollen on a flower head.





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