Pollen

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For other uses, see Pollen (disambiguation).
"Exine" redirects here. It is not to be confused with Exene.

Tulip anther with many grains of pollen

Closeup image of a cactus flower and its stamens

Scanning electron microscopeimage (500x magnification) 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).
Pollen is a fine to coarse powdery substance comprising pollen grains which are
male microgametophytes of seed plants, which produce male gametes (sperm cells). Pollen grains have a hard
coat made of sporopollenin that protects the gametophytes during the process of their movement from
the stamens to the pistil of flowering plants, or from the male cone to the female cone of coniferous plants. If
pollen lands on a compatible pistil or female cone, it germinates, producing a pollen tube that transfers
the sperm to the ovule containing the female gametophyte. Individual pollen grains are small enough to require
magnification to see detail. The study of pollen is called palynologyand is highly useful
in paleoecology, paleontology, archaeology, and forensics.
Pollen in plants is used for transferring haploid male genetic material from the anther of a single flower to the
stigma of another in cross-pollination.[1] In a case of self-pollination, this process takes place from the anther of
a flower to the stigma of the same flower.[1]

Contents

 1The structure and formation of pollen

o 1.1Formation
o 1.2Structure
 2Pollination
 3Pollen in the fossil record
 4Allergy to pollen
o 4.1Treatment
 5Nutrition
o 5.1In humans
o 5.2Parasites
 6Forensic palynology
 7Spiritual purposes
 8See also
 9References
 10Bibliography
 11External links

The structure and formation of pollen[edit]

Triporate pollen of Oenothera speciosa

Pollen of Lilium auratumshowing single sulcus (monosulcate)

Arabis pollen has three colpi and prominent surface structure.

Pollens/Microspores of Lycopersicon esculentum at coenocytic tetrad stage of development observed through oil
immersion microscope; the chromosomes of what will become four pollen grains can be seen.
Apple pollen under microscopy
Pollen itself is not the male gamete.[2] 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, and the generative cell divides to form the
two sperm cells.
Formation[edit]
Pollen is produced in the microsporangia in the male cone of a conifer or other gymnosperm or in the anthers
of an angiosperm flower. Pollen grains come in a wide variety of shapes, sizes, and surface markings
characteristic of the species (see electron micrograph, right). Pollen grains of pines, firs, and spruces are
winged. The smallest pollen grain, that of the forget-me-not (Myosotis spp.),[which?] is around 6 µm (0.006 mm)
in diameter.[citation needed] Wind-borne pollen grains can be as large as about 90–100 µm.[3]
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 prior
to cytokinesis, 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.[4]
During microgametogenesis, the unicellular microspores undergo mitosis and develop into
mature microgametophytes containing the gametes.[5] In some flowering plants,[which?] germination of the pollen
grain may begin even before it leaves the microsporangium, with the generative cell forming the two sperm
cells.
Structure[edit]
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 called
the endospore or intine, and a tough resistant outer cuticularized wall composed largely
of sporopollenincalled 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).
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.[6] They allow shrinking and swelling of the grain caused by changes in moisture content.
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.[7] 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.[8] However the term
inaperturate covers a wide range of morphological types, such as functionally inaperturate (cryptoaperturate)
and omniaperturate.[4] Inaperaturate pollen grains often have thin walls, which facilitates pollen
tube germination at any position.[6] Terms such as uniaperturate and triaperturate refer to the number of
apertures present (one and three respectively).
The orientation of furrows (relative to the original tetrad of microspores) classifies the pollen
as sulcateor colpate. Sulcate pollen has a furrow across the middle of what was the outer face when the pollen
grain was in its tetrad.[9] If the pollen has only a single sulcus, it is described as monosulcate, has two sulci,
as bisulcate, or more, as polysulcate.[10][11] Colpate pollen has furrows other than across the middle of the outer
faces.[9] Eudicots have pollen with three colpi (tricolpate) or with shapes that are evolutionarily derived from
tricolpate pollen.[12] The evolutionary trend in plants has been from monosulcate to polycolpate or polyporate
pollen.[9]
Additionally, gymnosperm pollen grains often have air bladders, or vesicles, called sacci. The sacci are not
actually balloons, but are sponge-like, and increase the buoyancy 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, spruce, and yellowwood trees all produce saccate pollen.[13]

Pollination[edit]
Main article: Pollination

European honey bee carrying pollen in a pollen basket back to the hive

Marmalade hoverfly, pollen on its face and legs, sitting on a rockrose.

Diadasia bee straddles flower carpels while visiting yellow Opuntia engelmannii cactus
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 mitesare specialized to feed on pollen,
and are called palynivores.
In non-flowering seed plants, pollen germinates in the pollen chamber, located beneath the micropyle,
underneath the integuments of the ovule. 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. Double-strand breaks in DNA that
arise during pollen tube 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.[14] However, the vegetative cell that is
responsible for tube elongation appears to lack this DNA repair capability.[14]

Pollen in the fossil record[edit]

Main article: Palynology
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. 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. Also, pollen analysis has
been widely used for reconstructing past changes in vegetation and their associated drivers [15]. Pollen is first
found in the fossil record in the late Devonianperiod,[16] [17] but at that time it is indistinguishable from
spores.[16] It increases in abundance until the present day.

Allergy to pollen[edit]
See also: Allergy season
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The examples and perspective in this article deal primarily with the United
States and do not represent a worldwide view of the subject. (September 2010)
This section needs additional citations for verification. (March 2013)
This section needs expansion with: information about allergies not in the nose,
e.g., skin reactions. You can help by adding to it. (March 2013)

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 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.[18]
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.[19]
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.
The number of people in the United States affected by hay fever is between 20 and 40 million,[20] and such
allergy has proven to be the most frequent allergic response in the nation. There are certain evidential
suggestions pointing out hay fever and similar allergies to be of hereditary origin. Individuals who suffer
from eczema or are asthmatic tend to be more susceptible to developing long-term hay fever.[21]
In Denmark, decades of rising temperatures cause pollen to appear earlier and in greater numbers, as well as
introduction of new species such as ragweed.[22]
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.[23]
Treatment[edit]
Main article: Allergic rhinitis § treatment
Antihistamines are effective at treating mild cases of pollinosis, this type of non-prescribed drugs
includes loratadine, cetirizine and chlorpheniramine. They do not prevent the discharge of histamine, 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 sprays.
Allergy immunotherapy (AIT) treatment involves administering doses of allergens to accustom the body to
pollen, thereby inducing specific long-term tolerance.[24] 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[edit]
Most major classes of predatory and parasitic arthropods contain species that eat pollen, despite the common
perception that bees are the primary pollen-consuming arthropod group. Many other Hymenoptera other than
bees consume pollen as adults, though only a small number feed on pollen as larvae (including
some ant larvae). Spiders are normally considered carnivores but pollen is an important source of food for
several species, particularly for spiderlings, 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.[citation
needed]
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 and Melyridae feed almost exclusively on pollen as adults, while various lineages within
larger families such as Curculionidae, Chrysomelidae, Cerambycidae, and Scarabaeidae are pollen specialists
even though most members of their families are not (e.g., only 36 of 40000 species of ground beetles, 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 1000 species). Similarly, Ladybird beetles mainly eat insects, but many
species also eat pollen, as either part or all of their diet. Hemiptera are mostly herbivores or omnivores but
pollen feeding is known (and has only been well studied in the Anthocoridae). 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).[25] Some species of fungus, including Fomes fomentarius, are able to break down
grains of pollen as a secondary nutrition source that is particularly high in nitrogen.[26] Pollen may be valuable
diet supplement for detritivores, providing them with nutrients needed for growth, development and
maturation.[27] 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.[27]
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.[28][29]
Although bats, butterflies and hummingbirds are not pollen eaters per se, their consumption of nectar in
flowers is an important aspect of the pollination process.
In humans[edit]
Bee pollen for human consumption is marketed as a food ingredient and as 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.[30]
Honey produced by bees from natural sources contains pollen derived p-coumaric acid,[31] an antioxidant and
natural bactericide that is also present in a wide variety of plants and plant-derived food products.[32]
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.[33]

Pine pollen (송화가루; Songhwa Garu) is traditionally consumed in Korea as an ingredient in sweets and
beverages[citation needed].
Parasites[edit]
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.[34] When this pollen has been tested for
parasites, it has been found that a multitude of pollinator viruses and eukaryotic parasites are present in the
pollen.[35][36] It is currently unclear if the parasites are introduced by the bee that collected the pollen or if it is
from contamination to the flower.[36][37] 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.[38] Several sterilization methods have been employed, though no method has been 100% effective at
sterilizing, without reducing the nutritional value, of the pollen [39]

Forensic palynology[edit]
Main article: Forensic palynology
An SEM micrograph of Redbudpollen. Scanning electron microscopes are major instruments in 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.[40] Pollen evidence can also reveal the season in which a particular object picked up the
pollen.[41] Pollen has been used to trace activity at mass graves in Bosnia,[42] catch a burglar who brushed
against a Hypericum bush during a crime,[43] and has even been proposed as an additive for bullets to enable
tracking them.[44]

Spiritual purposes[edit]
In some Native American religions, pollen is used in prayers and rituals to symbolize life and renewal
by sanctifying objects, dancing grounds, trails, and sandpaintings. It may also be sprinkled over heads or in
mouths. Many Navajo people believe the body becomes holy when it travels over a trail sprinkled with
pollen. [45]

See also[edit]
 European Pollen Database
 Evolution of sex
 Microsporangia
 Pollen calendar
 Pollen count
 Pollen DNA barcoding
 Pollen source
 Polyphenol antioxidant
 Palynology

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dePamphilis, Claude W.; Toth, Amy L.; Cox-Foster, Diana L.; Traveset, Anna
(22 December 2010). "RNA Viruses in Hymenopteran Pollinators: Evidence
of Inter-Taxa Virus Transmission via Pollen and Potential Impact on Non-
Apis Hymenopteran Species". PLoS ONE. 5 (12):
e14357. doi:10.1371/journal.pone.0014357. PMC 3008715. PMID 21203504
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 37. ^ Graystock, Peter; Goulson, Dave; Hughes, William O. H. (5 August
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Bibliography[edit]
 Davis, Owen (1999). "Palynology — Pollen". University of Arizona. Department
of Geosciences.
 Simpson, Michael G. (2011). Plant Systematics. Academic Press. ISBN 978-0-08-
051404-8. Retrieved 12 February 2014.
 Singh, Gurcharan (2004). Plant Systematics: An Integrated Approach. Science
Publishers. ISBN 978-1-57808-351-0. Retrieved 23 January 2014.

External links[edit]
Wikimedia Commons has
media related to Pollen.

Scholia has a topic profile

for Pollen.

 Pollen and Spore Identification Literature

 Pollen micrographs at SEM and confocal microscope
 The flight of a pollen cloud
  PalDat (database comprising palynological data from a variety of plant
families)
 Pollen-Wiki - A digital Pollen-Atlas, abgerufen am 09. Februar 2018.
 YouTube video of pollen clouds from Juncus gerardii plants

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