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Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male
gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce ...
s (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophytes during the process of their movement from the
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the fila ...
s to the pistil of flowering plants, or from the male cone to the female cone of
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, '' Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμν ...
s. If pollen lands on a compatible pistil or female cone, it germinates, producing a
pollen tube A pollen tube is a tubular structure produced by the male gametophyte of seed plants when it germinates. Pollen tube elongation is an integral stage in the plant life cycle. The pollen tube acts as a conduit to transport the male gamete cells fr ...
that transfers the
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, ...
to the
ovule In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the '' integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the ...
containing the female gametophyte. Individual pollen grains are small enough to require magnification to see detail. The study of pollen is called palynology and is highly useful in paleoecology,
paleontology Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
,
archaeology Archaeology or archeology is the scientific study of human activity through the recovery and analysis of material culture. The archaeological record consists of artifacts, architecture, biofacts or ecofacts, sites, and cultural landsc ...
, and
forensics Forensic science, also known as criminalistics, is the application of science to criminal and civil laws, mainly—on the criminal side—during criminal investigation, as governed by the legal standards of admissible evidence and crimin ...
. Pollen in plants is used for transferring haploid male genetic material from the
anther The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...
of a single flower to the stigma of another in cross-pollination. In a case of self-pollination, this process takes place from the anther of a flower to the stigma of the same flower. Pollen is infrequently used as food and food supplement. Because of agricultural practices, it is often contaminated by agricultural pesticides.


Structure and formation

Pollen itself is not the male gamete. It is a
gametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the ...
, something that could be considered an entire organism, which then produces the male gamete. 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. In flowering plants the vegetative tube cell produces the
pollen tube A pollen tube is a tubular structure produced by the male gametophyte of seed plants when it germinates. Pollen tube elongation is an integral stage in the plant life cycle. The pollen tube acts as a conduit to transport the male gamete cells fr ...
, and the generative cell divides to form the two sperm nuclei. File:Oenothera speciosa pollen 200x.jpg, Triporate pollen of '' Oenothera speciosa'' File:Lilium auratum - pollen.jpg, Pollen of '' Lilium auratum'' showing single sulcus (monosulcate) File:Arabis voch1-4.jpg, '' Arabis'' pollen has three colpi and prominent surface structure. File:Жизнеспособность пыльцы 13.jpg, Apple pollen under microscopy File:Fire lily pollens on an insect's hair.jpg, Pollen of Lilium bulbiferum on an insect's hair under microscope


Formation

Pollen is produced in the microsporangia in the male cone of a conifer or other
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, '' Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμν ...
or in the anthers of an
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants ...
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanis ...
. Pollen grains come in a wide variety of shapes, sizes, and surface markings characteristic of the species (see electron micrograph, right). Pollen grains of
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanic ...
s, firs, and
spruce A spruce is a tree of the genus ''Picea'' (), a genus of about 35 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal (taiga) regions of the Earth. ''Picea'' is the sole genus in the subfam ...
s are winged. The smallest pollen grain, that of the forget-me-not (''Myosotis'' spp.), is 2.5-5 
µm The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer ( American spelling), also commonly known as a micron, is a unit of length in the International System of Uni ...
(0.005 mm) in diameter. Corn pollen grains are large, about 90–100 µm. Most grass pollen is around 20-25 µm. 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 within 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. In a process called microsporogenesis, four haploid microspores are produced from each diploid sporogenous cell (microsporocyte, pollen mother cell or meiocyte), 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. Two basic types of microsporogenesis are recognised, simultaneous and successive. In simultaneous microsporogenesis meiotic steps I and II are completed before
cytokinesis Cytokinesis () is the part of the cell division process during which the cytoplasm of a single eukaryotic cell divides into two daughter cells. Cytoplasmic division begins during or after the late stages of nuclear division in mitosis and mei ...
, whereas in successive microsporogenesis cytokinesis follows. While there may be a continuum with intermediate forms, the type of microsporogenesis has systematic significance. The predominant form amongst the monocots is successive, but there are important exceptions. During microgametogenesis, the unicellular microspores undergo mitosis and develop into mature microgametophytes containing the gametes. In some flowering plants,
germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fe ...
of the pollen grain may begin even before it leaves the microsporangium, with the generative cell forming the two sperm cells.


Structure

Except in the case of some submerged aquatic plants, the mature pollen grain has a double wall. The vegetative and generative cells are surrounded by a thin delicate wall of unaltered
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wa ...
called the endospore or intine, and a tough resistant outer cuticularized wall composed largely of sporopollenin called the 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. The spines may be less than a micron in length (spinulus, plural spinuli) referred to as spinulose (scabrate), or longer than a micron (echina, echinae) referred to as echinate. Various terms also describe the sculpturing such as reticulate, a net like appearance consisting of elements (murus, muri) separated from each other by a lumen (plural lumina). These reticulations may also be referred to as brochi. The pollen wall protects the sperm 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, which prevents the pollen grain from shrinking and crushing the genetic material during desiccation, 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. Pollen apertures are regions of the pollen wall that may involve exine thinning or a significant reduction in exine thickness. They allow shrinking and swelling of the grain caused by changes in moisture content. The process of shrinking the grain is called harmomegathy. Elongated apertures or furrows in the pollen grain are called colpi (singular: colpus) or sulci (singular: sulcus). Apertures that are more circular are called pores. Colpi, sulci and pores are major features in the identification of classes of pollen. Pollen may be referred to as inaperturate (apertures absent) or aperturate (apertures present). The aperture may have a lid ( operculum), hence is described as operculate. However the term inaperturate covers a wide range of morphological types, such as functionally inaperturate (cryptoaperturate) and omniaperturate. Inaperaturate pollen grains often have thin walls, which facilitates
pollen tube A pollen tube is a tubular structure produced by the male gametophyte of seed plants when it germinates. Pollen tube elongation is an integral stage in the plant life cycle. The pollen tube acts as a conduit to transport the male gamete cells fr ...
germination at any position. Terms such as uniaperturate and triaperturate refer to the number of apertures present (one and three respectively). Spiraperturate refers to one or more apertures being spirally shaped. The orientation of furrows (relative to the original tetrad of microspores) classifies the pollen as sulcate or colpate. Sulcate pollen has a furrow across the middle of what was the outer face when the pollen grain was in its tetrad. If the pollen has only a single sulcus, it is described as monosulcate, has two sulci, as bisulcate, or more, as polysulcate. Colpate pollen has furrows other than across the middle of the outer faces, and similarly may be described as polycolpate if more than two. Syncolpate pollen grains have two or more colpi that are fused at the ends.
Eudicots The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons. Traditionally they were called tricolpates or non-magnoliid dicot ...
have pollen with three colpi (
tricolpate The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons. Traditionally they were called tricolpates or non-magnoliid dicots ...
) or with shapes that are evolutionarily derived from tricolpate pollen. The evolutionary trend in plants has been from monosulcate to polycolpate or polyporate pollen. Additionally,
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, '' Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμν ...
pollen grains often have air bladders, or vesicles, called sacci. The sacci are not actually balloons, but are sponge-like, and increase the
buoyancy Buoyancy (), or upthrust, is an upward force exerted by a fluid that opposes the weight of a partially or fully immersed object. In a column of fluid, pressure increases with depth as a result of the weight of the overlying fluid. Thus the ...
of the pollen grain and help keep it aloft in the wind, as most gymnosperms are anemophilous. Pollen can be monosaccate, (containing one saccus) or bisaccate (containing two sacci). Modern
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanic ...
,
spruce A spruce is a tree of the genus ''Picea'' (), a genus of about 35 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal (taiga) regions of the Earth. ''Picea'' is the sole genus in the subfam ...
, and yellowwood trees all produce saccate pollen.


Pollination

The transfer of pollen grains to the female reproductive structure ('' pistil'' in angiosperms) is called
pollination Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...
. 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 Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
-rich, for dispersal by
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the m ...
s attracted to their flowers. Many insects and some
mite Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear e ...
s are specialized to feed on pollen, and are called
palynivore In zoology, a palynivore /pəˈlɪnəvɔːɹ/, meaning "pollen eater" (from Greek παλύνω palunō, "strew, sprinkle", and Latin, vorare, meaning "to devour") is an herbivorous animal which selectively eats the nutrient-rich pollen produce ...
s. In non-flowering seed plants, pollen germinates in the pollen chamber, located beneath the
micropyle Micropyle may refer to: * Micropyle (botany) a minute opening in the integument of an ovule of a seed plant. * Micropyle (zoology) A micropyle is a pore in the membrane covering the ovum, through which a sperm enters. Micropyles are also found in s ...
, underneath the integuments of the ovule. A
pollen tube A pollen tube is a tubular structure produced by the male gametophyte of seed plants when it germinates. Pollen tube elongation is an integral stage in the plant life cycle. The pollen tube acts as a conduit to transport the male gamete cells fr ...
is produced, which grows into the
nucellus In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the '' nucellus'' (or remnant of the megasporangium), and the ...
to provide nutrients for the developing sperm cells. Sperm cells of
Pinophyta Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ext ...
and
Gnetophyta Gnetophyta () is a division of plants (alternatively considered the subclass Gnetidae or order Gnetales), grouped within the gymnosperms (which also includes conifers, cycads, and ginkgos), that consists of some 70 species across the three ...
are without
flagella A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have fro ...
, and are carried by the pollen tube, while those of
Cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male o ...
ophyta 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 A pollen tube is a tubular structure produced by the male gametophyte of seed plants when it germinates. Pollen tube elongation is an integral stage in the plant life cycle. The pollen tube acts as a conduit to transport the male gamete cells fr ...
, which grows down the tissue of the style to the
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
, and makes its way along the
placenta The placenta is a temporary embryonic and later fetal organ that begins developing from the blastocyst shortly after implantation. It plays critical roles in facilitating nutrient, gas and waste exchange between the physically separate mate ...
, guided by projections or hairs, to the micropyle of an
ovule In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the '' integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the ...
. 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 A pollen tube is a tubular structure produced by the male gametophyte of seed plants when it germinates. Pollen tube elongation is an integral stage in the plant life cycle. The pollen tube acts as a conduit to transport the male gamete cells fr ...
. Double-strand breaks in DNA that arise during
pollen tube A pollen tube is a tubular structure produced by the male gametophyte of seed plants when it germinates. Pollen tube elongation is an integral stage in the plant life cycle. The pollen tube acts as a conduit to transport the male gamete cells fr ...
growth appear to be efficiently repaired in the generative cell that carries the male genomic information to be passed on to the next plant generation. However, the vegetative cell that is responsible for tube elongation appears to lack this
DNA repair DNA repair is a collection of processes by which a cell identifies and corrects damage to the DNA molecules that encode its genome. In human cells, both normal metabolic activities and environmental factors such as radiation can cause DNA d ...
capability.


In the fossil record

The sporopollenin outer sheath of pollen grains affords them some resistance to the rigours of the fossilisation process that destroy weaker objects; it is also produced in huge quantities. 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 Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictionary of ...
and to gain information about the abundance and variety of plants alive — which can itself yield important information about paleoclimates. Also, pollen analysis has been widely used for reconstructing past changes in vegetation and their associated drivers. Pollen is first found in the
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
record in the late
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
period, but at that time it is indistinguishable from spores. It increases in abundance until the present day.


Allergy to pollen

Nasal allergy to pollen is called pollinosis, and allergy specifically to grass 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. Pollen allergies are common in polar and temperate climate zones, where production of pollen is seasonal. In the tropics pollen production varies less by the season, and allergic reactions less. In northern Europe, common pollens for allergies are those of
birch A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech- oak family Fagaceae. The genus ''Betula'' cont ...
and alder, and in late summer wormwood and different forms of hay. Grass pollen is also associated with asthma exacerbations in some people, a phenomenon termed thunderstorm asthma. 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 Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a state in the Southwestern United States. It is the 6th largest and the 14th most populous of the 50 states. Its capital and largest city is Phoenix. Arizona is part of the Fou ...
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 A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech- oak family Fagaceae. The genus ''Betula'' cont ...
,
hickory Hickory is a common name for trees composing the genus ''Carya'', which includes around 18 species. Five or six species are native to China, Indochina, and India (Assam), as many as twelve are native to the United States, four are found in Mex ...
, pecan, and early summer
grass Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns a ...
es may also induce pollen allergies. Most cultivated plants with showy flowers are entomophilous and do not cause pollen allergies. Symptoms of pollen allergy include
sneezing A sneeze (also known as sternutation) is a semi-autonomous, convulsive expulsion of air from the lungs through the nose and mouth, usually caused by foreign particles irritating the nasal mucosa. A sneeze expels air forcibly from the mouth an ...
, itchy, or runny nose, nasal congestion, red, itchy, and watery eyes. Substances, including pollen, that cause allergies can trigger asthma. A study found a 54% increased chance of
asthma Asthma is a long-term inflammatory disease of the airways of the lungs. It is characterized by variable and recurring symptoms, reversible airflow obstruction, and easily triggered bronchospasms. Symptoms include episodes of wheezing, co ...
attacks when exposed to pollen. The number of people in the United States affected by hay fever is between 20 and 40 million, including around 6.1 million children and such allergy has proven to be the most frequent
allergic response An allergic response is a hypersensitive immune reaction to a substance that normally is harmless or would not cause an immune response in everyone. An allergic response may cause harmful symptoms such as itching or inflammation or tissue inju ...
in the nation. Hay fever affects about 20% of Canadians and the prevalence is increasing. There are certain evidential suggestions pointing out hay fever and similar allergies to be of hereditary origin. Individuals who suffer from
eczema Dermatitis is inflammation of the skin, typically characterized by itchiness, redness and a rash. In cases of short duration, there may be small blisters, while in long-term cases the skin may become thickened. The area of skin involved c ...
or are
asthma Asthma is a long-term inflammatory disease of the airways of the lungs. It is characterized by variable and recurring symptoms, reversible airflow obstruction, and easily triggered bronchospasms. Symptoms include episodes of wheezing, co ...
tic tend to be more susceptible to developing long-term hay fever. Since 1990, pollen seasons have gotten longer and more pollen-filled, and climate change is responsible, according to a new study. The researchers attributed roughly half of the lengthening pollen seasons and 8% of the trend in pollen concentrations to climate changes driven by human activity. In
Denmark ) , song = ( en, "King Christian stood by the lofty mast") , song_type = National and royal anthem , image_map = EU-Denmark.svg , map_caption = , subdivision_type = Sovereign state , subdivision_name = Kingdom of Denmark , establish ...
, decades of rising temperatures cause pollen to appear earlier and in greater numbers, as well as introduction of new species such as ragweed. The most efficient way to handle a pollen allergy is by preventing contact with the material. Individuals carrying the ailment may at first believe that they have a simple summer cold, but hay fever becomes more evident when the apparent cold does not disappear. The confirmation of hay fever can be obtained after examination by a general physician.


Treatment

Antihistamines are effective at treating mild cases of pollinosis; this type of non-prescribed drugs includes loratadine, cetirizine and
chlorpheniramine Chlorphenamine (CP, CPM), also known as chlorpheniramine, is an antihistamine used to treat the symptoms of allergic conditions such as allergic rhinitis (hay fever). It is taken by mouth. The medication takes effect within two hours and lasts ...
. They do not prevent the discharge of
histamine Histamine is an organic nitrogenous compound involved in local immune responses, as well as regulating physiological functions in the gut and acting as a neurotransmitter for the brain, spinal cord, and uterus. Since histamine was discover ...
, but it has been proven that they do prevent a part of the chain reaction activated by this biogenic amine, which considerably lowers hay fever symptoms. Decongestants can be administered in different ways such as tablets and
nasal spray Nasal sprays are used to deliver medications locally in the nasal cavities or systemically. They are used locally for conditions such as nasal congestion and allergic rhinitis. In some situations, the nasal delivery route is preferred for syst ...
s. Allergy immunotherapy (AIT) treatment involves administering doses of allergens to accustom the body to pollen, thereby inducing specific long-term tolerance. Allergy immunotherapy can be administered orally (as sublingual tablets or sublingual drops), or by injections under the skin (subcutaneous). Discovered by Leonard Noon and John Freeman in 1911, allergy immunotherapy represents the only causative treatment for respiratory allergies.


Nutrition

Most major classes of predatory and parasitic
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s contain species that eat pollen, despite the common perception that
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfami ...
s are the primary pollen-consuming arthropod group. Many
Hymenoptera Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic. Females typic ...
other than bees consume pollen as adults, though only a small number feed on pollen as
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e (including some ant larvae).
Spider Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s are normally considered carnivores but pollen is an important source of food for several species, particularly for
spider Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
lings, which catch pollen on their webs. It is not clear how spiderlings manage to eat pollen however, since their mouths are not large enough to consume pollen grains. Some predatory mites also feed on pollen, with some species being able to subsist solely on pollen, such as '' Euseius tularensis'', which feeds on the pollen of dozens of plant species. Members of some beetle families such as
Mordellidae The Mordellidae are a family of beetles commonly known as tumbling flower beetles for the typical irregular movements they make when escaping predators, or as pintail beetles due to their abdominal tip which aids them in performing these tumbling ...
and Melyridae feed almost exclusively on pollen as adults, while various lineages within larger families such as
Curculionidae The Curculionidae are a family of weevils, commonly called snout beetles or true weevils. They are one of the largest animal families, with 6,800 genera and 83,000 species described worldwide. They are the sister group to the family Brentidae. ...
,
Chrysomelidae The insects of the beetle family Chrysomelidae are commonly known as leaf beetles, and include over 37,000 (and probably at least 50,000) species in more than 2,500 genera, making up one of the largest and most commonly encountered of all beetle ...
, Cerambycidae, and Scarabaeidae are pollen specialists even though most members of their families are not (e.g., only 36 of 40,000 species of
ground beetle Ground beetles are a large, cosmopolitan family of beetles, the Carabidae, with more than 40,000 species worldwide, around 2,000 of which are found in North America and 2,700 in Europe. As of 2015, it is one of the 10 most species-rich animal f ...
s, which are typically predatory, have been shown to eat pollen—but this is thought to be a severe underestimate as the feeding habits are only known for 1,000 species). Similarly, Ladybird beetles mainly eat insects, but many species also eat pollen, as either part or all of their diet. Hemiptera are mostly
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
s or
omnivore An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nu ...
s but pollen feeding is known (and has only been well studied in the
Anthocoridae Anthocoridae is a family of bugs, commonly called minute pirate bugs or flower bugs. Worldwide there are 500-600 species. Description Anthocoridae are 1.5–5 mm long and have soft, elongated oval, flat bodies, often patterned in black an ...
). Many adult flies, especially Syrphidae, feed on pollen, and three UK syrphid species feed strictly on pollen (syrphids, like all flies, cannot eat pollen directly due to the structure of their mouthparts, but can consume pollen contents that are dissolved in a fluid). Some species of fungus, including ''
Fomes fomentarius ''Fomes fomentarius'' (commonly known as the tinder fungus, false tinder fungus, hoof fungus, tinder conk, tinder polypore or ice man fungus) is a species of fungal plant pathogen found in Europe, Asia, Africa and North America. The species produ ...
'', are able to break down grains of pollen as a secondary nutrition source that is particularly high in nitrogen. Pollen may be valuable diet supplement for
detritivore Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrate ...
s, providing them with nutrients needed for growth, development and maturation. It was suggested that obtaining nutrients from pollen, deposited on the forest floor during periods of pollen rains, allows fungi to decompose nutritionally scarce litter. Some species of '' Heliconius'' butterflies consume pollen as adults, which appears to be a valuable nutrient source, and these species are more distasteful to predators than the non-pollen consuming species. Although bats,
butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises ...
and
hummingbird Hummingbirds are birds native to the Americas and comprise the Family (biology), biological family Trochilidae. With about 361 species and 113 genus, genera, they occur from Alaska to Tierra del Fuego, but the vast majority of the species are ...
s are not pollen eaters ''
per se Per se may refer to: * ''per se'', a Latin phrase meaning "by itself" or "in itself". * Illegal ''per se'', the legal usage in criminal and antitrust law * Negligence ''per se'', legal use in tort law *Per Se (restaurant), a New York City restauran ...
'', their consumption of
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualist ...
in flowers is an important aspect of the
pollination Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...
process.


In humans

Bee pollen for human consumption is marketed as a
food Food is any substance consumed by an organism for nutritional support. Food is usually of plant, animal, or fungal origin, and contains essential nutrients, such as carbohydrates, fats, proteins, vitamins, or minerals. The substance is in ...
ingredient and as a
dietary supplement A dietary supplement is a manufactured product intended to supplement one's diet by taking a pill, capsule, tablet, powder, or liquid. A supplement can provide nutrients either extracted from food sources or that are synthetic in order ...
. The largest constituent is
carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may o ...
s, with protein content ranging from 7 to 35 percent depending on the plant species collected by bees.
Honey Honey is a sweet and viscous substance made by several bees, the best-known of which are honey bees. Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of plants (primar ...
produced by bees from natural sources contains pollen derived p-coumaric acid, an
antioxidant Antioxidants are compounds that inhibit oxidation, a chemical reaction that can produce free radicals. This can lead to polymerization and other chain reactions. They are frequently added to industrial products, such as fuels and lubrica ...
and natural
bactericide A bactericide or bacteriocide, sometimes abbreviated Bcidal, is a substance which kills bacteria. Bactericides are disinfectants, antiseptics, or antibiotics. However, material surfaces can also have bactericidal properties based solely on their ...
that is also present in a wide variety of plants and plant-derived food products. The U.S. Food and Drug Administration (FDA) has not found any harmful effects of bee pollen consumption, except from the usual allergies. However, FDA does not allow bee pollen marketers in the United States to make health claims about their produce, as no scientific basis for these has ever been proven. 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. Pine pollen () is traditionally consumed in Korea as an ingredient in sweets and beverages.


Parasites

The growing industries in pollen harvesting for human and bee consumption rely on harvesting pollen baskets from honey bees as they return to their hives using a ''pollen trap''. When this pollen has been tested for parasites, it has been found that a multitude of viruses and eukaryotic parasites are present in the pollen. It is currently unclear if the parasites are introduced by the bee that collected the pollen or if it is from the flower. Though this is not likely to pose a risk to humans, it is a major issue for the bumblebee rearing industry that relies on thousands of tonnes of honey bee collected pollen per year. Several sterilization methods have been employed, though no method has been 100% effective at sterilisation without reducing the nutritional value of the pollen


Forensic palynology

In
forensic biology Forensic biology is the application of biology to associate a person(s), whether suspect or victim, to a location, an item (or collection of items), another person (victim or suspect, respectively). It can be utilized to further investigations for ...
, 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 Bosnia, 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.


Spiritual purposes

In some Native American religions, pollen was used in
prayer Prayer is an invocation or act that seeks to activate a rapport with an object of worship through deliberate communication. In the narrow sense, the term refers to an act of supplication or intercession directed towards a deity or a deifie ...
s and rituals to symbolize life and renewal by
sanctifying Sanctification (or in its verb form, sanctify) literally means "to set apart for special use or purpose", that is, to make holy or sacred (compare la, sanctus). Therefore, sanctification refers to the state or process of being set apart, i.e. " ...
objects, dancing grounds, trails, and sandpaintings. It may also be sprinkled over heads or in mouths. Many
Navajo The Navajo (; British English: Navaho; nv, Diné or ') are a Native Americans in the United States, Native American people of the Southwestern United States. With more than 399,494 enrolled tribal members , the Navajo Nation is the largest fe ...
people believed the body became holy when it traveled over a trail sprinkled with pollen.


Pollen grain staining

For agricultural research purposes, assessing the viability of pollen grains can be necessary and illuminating. A very common, efficient method to do so is known as Alexander's stain. This differential stain consists of
ethanol Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an alcohol with the chemical formula . Its formula can be also written as or (an ethyl group linked to a ...
, malachite green, distilled water,
glycerol Glycerol (), also called glycerine in British English and glycerin in American English, is a simple triol compound. It is a colorless, odorless, viscous liquid that is sweet-tasting and non-toxic. The glycerol backbone is found in lipids known ...
,
phenol Phenol (also called carbolic acid) is an aromatic organic compound with the molecular formula . It is a white crystalline solid that is volatile. The molecule consists of a phenyl group () bonded to a hydroxy group (). Mildly acidic, it r ...
, chloral hydrate,
acid fuchsin Acid fuchsin or fuchsine acid, (also called Acid Violet 19 and C.I. 42685) is an acidic magenta dye with the chemical formula C20H17N3Na2O9S3. It is a sodium sulfonate derivative of fuchsine. Acid fuchsin has wide use in histology, and is one of ...
,
orange G Orange G also called C.I. 16230, Acid Orange 10, or orange gelb is a synthetic azo dye used in histology in many staining formulations. It usually comes as a disodium salt. It has the appearance of orange crystals or powder. Staining Orange G ...
, and glacial acetic acid. In angiosperms and gymnosperms non-aborted pollen grain will appear red or pink, and aborted pollen grains will appear blue or slightly green.


See also

* European Pollen Database * Evolution of sex * Honeybee starvation * Pollen calendar *
Pollen count A pollen count is the measurement of the number of grains of pollen in a cubic meter of air. High pollen counts can sometimes lead to increased rates of allergic reaction for those with allergic disorders. Usually, the counts are announced for spe ...
* Pollen DNA barcoding *
Pollen source The term pollen source is often used in the context of beekeeping and refers to flowering plants as a source of pollen for bees or other insects. Bees collect pollen as a protein source to raise their brood. For the plant, the pollinizer, thi ...
*
Polyphenol antioxidant A polyphenol antioxidant is a hypothetical type of antioxidant containing a polyphenolic substructure and studied in vitro. Numbering over 4,000 distinct species mostly from plants, polyphenols may have antioxidant activity in vitro, but are unlik ...


References


Bibliography

* * * * * * *


External links


Pollen and Spore Identification Literature





PalDat (database comprising palynological data from a variety of plant families)

''Pollen-Wiki - A digital Pollen-Atlas''
retrieved 9 February 2018.
YouTube video of pollen clouds from Juncus gerardii plants

Information on Pollen Count & Pollen Allergies

Important Facts About Pollen Counts
{{Authority control Plant anatomy Plant morphology Pollination Allergology