Pollen is a fine to coarse powder consisting of microgametophytes (pollen grains), which produce the male gametes (sperm cells) of seed plants. A hard coat covering the pollen grain protects the sperm cells during the process of their movement between the stamens of the flower to the pistil of the next flower.
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, sizes, and surface markings characteristic of the species (see Electron micrograph at top right). Most, but certainly not all, are spherical. 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 microsporocytes, 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 made up of a resistant compound called sporopollenin; the intine is made up of cellulose and pectin. The exine is what is preserved in the fossil record.
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.
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.
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”.
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.
Arizona 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.
A variety of producers have started selling pollen for human consumption, often marketed as a healthy food. Like many other foods, pollen contains water, amino acids, proteins, lipids, carbohydrates, minerals, vitamins, enzymes and other micronutrients. Composition depends on plant species collected by bees. Supplements of pollen have been used in a number of treatments, e.g. treatment of anemia, disorders of digestive system, mental disorders like depression, asthenia and alcohol dependence.
The 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.