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Pollen

For other uses, see Pollen (disambiguation).


Exine redirects here. It is not to be confused with
Exene.
Pollen is a ne to coarse powdery substance comprising

Scanning electron microscope image of pollen grains from a


variety of common plants: sunower (Helianthus annuus),
morning glory (Ipomoea purpurea), prairie hollyhock (Sidalcea
malviora), oriental lily (Lilium auratum), evening primrose
(Oenothera fruticosa), and castor bean (Ricinus communis).

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 owering 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 magnication to see detail. The study of pollen is
called palynology and is highly useful in paleoecology,
paleontology, archaeology, and forensics.

Tulip anther with many grains of pollen

Pollen in plants is used for transferring haploid male genetic material from the anther of a single ower to the
stigma of another in cross-pollination. In a case of selfpollination, this process takes place from the anther of a
ower to the stigma of the same ower.

1 The structure and formation of


pollen
Pollen itself is not the male gamete.[1] Each pollen grain
contains vegetative (non-reproductive) cells (only a single cell in most owering plants but several in other seed
plants) and a generative (reproductive) cell. In owering
plants the vegetative tube cell produces the pollen tube,
and the generative cell divides to form the two sperm

Closeup image of a cactus ower and its stamens

pollen grains which are male microgametophytes of seed


1

1 THE STRUCTURE AND FORMATION OF POLLEN

Triporate pollen of Oenothera speciosa

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.

Pollen of Lilium auratum showing single sulcus (monosulcate)

Apple pollen under microscopy

Arabis pollen has three colpi and prominent surface structure.

cells.

1.1

Formation

Pollen is produced in the 'microsporangium' (contained in


the anther of an angiosperm ower, male cone of a coniferous plant, or male cone of other seed plants). Pollen
grains come in a wide variety of shapes (most often spher-

ical), sizes, and surface markings characteristic of the


species (see electron micrograph, right). Pollen grains of
pines, rs, and spruces are winged. The smallest pollen
grain, that of the forget-me-not (Myosotis spp.), is around
6 m (0.006 mm) in diameter. Wind-borne pollen grains
can be as large as about 90100 m.[2]
In angiosperms, during ower development the anther is
composed of a mass of cells that appear undierentiated,
except for a partially dierentiated dermis. As the ower
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.

3
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 signicance.
The predominant form amongst the monocots is successive, but there are important exceptions.[3]

called sporopollenin.
Pollen apertures are regions of the pollen wall that may
involve exine thinning or a signicant reduction in exine thickness.[5] 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 identication
of classes of pollen.[6] 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.[7] However the term
inaperturate covers a wide range of morphological types,
such as functionally inaperturate (cryptoaperturate) and
omniaperturate.[3] Inaperaturate pollen grains often have
thin walls, which facilitates pollen tube germination at any
position.[5]

The orientation of furrows (relative to the original tetrad


of microspores) classies 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.[8]
If the pollen has only a single sulcus, it is described as
monosulcate.[9][10] Colpate pollen has furrows other than
across the middle of the outer faces.[8] Eudicots have
pollen with three colpi (tricolpate) or with shapes that
are evolutionarily derived from tricolpate pollen.[11] The
1.2 Structure
evolutionary trend in plants has been from monosulcate
[8]
Except in the case of some submerged aquatic plants, the to polycolpate or polyporate pollen.
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 2 Pollination
tough resistant outer cuticularized wall composed largely
of sporopollenin called the exospore or exine. The exine Main article: Pollination
often bears spines or warts, or is variously sculptured, and
The transfer of pollen grains to the female reproducthe 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).
In the microgametogenesis, the unicellular microspores undergoes mitosis and develops into mature
microgametophytes containing the gametes.[4] In some
owering plants, germination of the pollen grain often
begins before it leaves the microsporangium, with the
generative cell forming the two sperm cells.

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

European honey bee carrying pollen in a pollen basket back to


the hive

tive 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-owering seed plants

ALLERGY TO POLLEN

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 pollentube.

3 Pollen in the fossil record


Main article: Palynology

Marmalade hovery, pollen on its face and legs, sitting on a


rockrose.

Pollens sporopollenin outer sheath aords 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. Pollen is rst found in the fossil record in the
late Devonian period and increases in abundance until the
present day.

4 Allergy to pollen
See also: Allergy season

Africanized honey bees visiting yellow Opuntia engelmannii


cactus

(e.g. pine trees) are characteristically anemophilous.


Anemophilous owering plants generally have inconspicuous owers. Entomophilous (literally insect-loving)
plants produce pollen that is relatively heavy, sticky and
protein-rich, for dispersal by insect pollinators attracted
to their owers. Many insects and some mites are specialized to feed on pollen, and are called palynivores.

Nasal allergy to pollen is called pollinosis, and allergy


specically 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 ower 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 ower for allergies. Since this plant is entomophilous (its pollen is dispersed by animals), its heavy,
sticky pollen does not become independently airborne.
In non-owering seed plants, pollen germinates in the Most late summer and fall pollen allergies are probably
[12]
pollen chamber, located beneath the micropyle, under- caused by ragweed, a widespread anemophilous plant.
neath the integuments of the ovule. A pollen tube is Arizona was once regarded as a haven for people with
produced, which grows into the nucellus to provide nu- pollen allergies, although several ragweed species grow in
trients for the developing sperm cells. Sperm cells of the desert. However, as suburbs grew and people began
Pinophyta and Gnetophyta are without agella, and are establishing irrigated lawns and gardens, more irritating
carried by the pollen tube, while those of Cycadophyta species of ragweed gained a foothold and Arizona lost its
and Ginkgophyta have many agella.
claim of freedom from hay fever.
When placed on the stigma of a owering plant, under fa- Anemophilous spring blooming plants such as oak, birch,
vorable circumstances, a pollen grain puts forth a pollen hickory, pecan, and early summer grasses may also intube, which grows down the tissue of the style to the duce pollen allergies. Most cultivated plants with showy

5.1

In humans

owers are entomophilous and do not cause pollen aller- an important source of food for several species, particgies.
ularly for spiderlings, which catch pollen on their webs.
The number of people in the United States aected by It is not clear how spiderlings manage to eat pollen howhay fever is between 20 and 40 million,[13] and such ever, since their mouths are not large enough to consume
allergy has proven to be the most frequent allergic re- pollen grains. Some predatory mites also feed on pollen,
sponse in the nation. There are certain evidential sugges- with some species being able to subsist solely on pollen,
tions pointing out hay fever and similar allergies to be of such as Euseius tularensis, which feeds on the pollen of
hereditary origin. Individuals who suer from eczema or dozens of plant species. Members of some beetle families such as Mordellidae and Melyridae feed almost exare asthmatic tend to be more susceptible to developing
clusively
on pollen as adults, while various lineages within
[14]
long-term hay fever.
larger families such as Curculionidae, Chrysomelidae,
In Denmark, decades of rising temperatures cause pollen Cerambycidae, and Scarabaeidae are pollen specialists
to appear earlier and in greater numbers, as well as intro- even though most members of their families are not (e.g.,
duction of new species such as ragweed.[15]
only 36 of 40000 species of ground beetles, which are
typically
predatory, have been shown to eat pollen
The most ecient way to handle a pollen allergy is by
but
this
is
thought to be a severe underestimate as the
preventing contact with the material. Individuals carrying
feeding
habits
are only known for 1000 species). Simthe ailment may at rst believe that they have a simple
Ladybird
beetles mainly eat insects, but many
ilarly,
summer cold, but hay fever becomes more evident when
species
also
eat
pollen,
as either part or all of their diet.
the apparent cold does not disappear. The conrmation of
Hemiptera
are
mostly
herbivores
or omnivores but pollen
hay fever can be obtained after examination by a general
[16]
feeding
is
known
(and
has
only
been
well studied in the
physician.
Anthocoridae). Many adult ies, especially Syrphidae,
feed on pollen, and three UK syrphid species feed strictly
on pollen (syrphids, like all ies, cannot eat pollen di4.1 Treatment
rectly due to the structure of their mouthparts, but can
consume pollen contents that are dissolved in a uid).[18]
Main article: Allergic rhinitis treatment
Some species of fungus, including Fomes fomentarius, are
able to break down grains of pollen as a secondary nutriAntihistamines are eective at treating mild cases of tion source that is particularly high in nitrogen.[19]
pollinosis, this type of non-prescribed drugs includes
loratadine, cetirizine and chlorphenamine. They do not Some species of Heliconius butteries consume pollen as
prevent the discharge of histamine, but it has been proven adults, which appears to be a valuable nutrient source,
to predators than
that they do prevent a part of the chain reaction activated and these species are more distasteful
[20][21]
the
non-pollen
consuming
species.
by this biogenic amine, which considerably lowers hay
fever symptoms.
Although bats, butteries and hummingbirds are not
pollen
eaters per se, their consumption of nectar in owDecongestants can be administered in dierent ways such
ers
is
an
important aspect of the pollination process.
as tablets and nasal sprays.
Allergy immunotherapy (AIT) treatment involves administering doses of allergens to accustom the body to pollen,
thereby inducing specic long-term tolerance.[17] 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 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

5.1 In humans
A variety of producers have started selling bee 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.[22]
Honey produced by bees from natural sources contains
pollen derived p-coumaric acid, an antioxidant.[23]
The U.S. Food and Drug Administration (FDA) has not
found any harmful eects 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 scientic 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

manufacturerss claim that pollen collecting helps the bee


colonies is also controversial.[24]
Pine pollen (
, Songhwa Garu) is traditionally consumed in Korea as an ingredient in sweets and beverages.

5.2

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.[25] 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.[26][27] It is
currently unclear if the parasites are introduced by the bee
that collected the pollen or if it is from contamination to
the ower.[27][28] 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.[29] Several sterilization methods
have been employed, though no method has been 100%
eective at steralizing, without reducing the nutritional
value, of the pollen [30]

Forensic palynology

Main article: Forensic palynology


In forensic biology, pollen can tell a lot about where

REFERENCES

7 See also
European Pollen Database
Evolution of sex
Microsporangia
Pollen calendar
Pollen count
Pollen source
Polyphenol antioxidant
Palynology

8 References
[1] Johnstone, Adam (2001). Biology: facts & practice for
A level. Oxford University Press. p. 95. ISBN 0-19914766-3.
[2] Pleasants, J. M.; Hellmich, R. L.; Dively, G. P.; Sears,
M. K.; Stanley-Horn, D. E.; Mattila, H. R.; Foster, J.
E.; Clark, P.; Jones, G. D. (2001). Corn pollen deposition on milkweeds in and near cornelds (Free full
text). Proceedings of the National Academy of Sciences
of the United States of America. 98 (21): 1191924.
doi:10.1073/pnas.211287498. PMC 59743 . PMID
11559840.
[3] Furness & Rudall 2001.
[4] Pollen Development University of Leicester
[5] Furness, Carol A.; Rudall, Paula J. (2004-03-01). Pollen
aperture evolution--a crucial factor for eudicot success?". Trends in Plant Science. 9 (3): 154158.
doi:10.1016/j.tplants.2004.01.001. ISSN 1360-1385.
PMID 15003239.
[6] Davis, Owen. Aperture. geo.arizona.edu.
[7] Furness & Rudall 2003.

An SEM micrograph of Redbud pollen. Scanning electron microscopes are major instruments in palynology.

[8] Sporne, Kenneth R. (1972). Some Observations on


the Evolution of Pollen Types in Dicotyledons. New
Phytologist. 71 (1): 181185. doi:10.1111/j.14698137.1972.tb04826.x.
[9] Simpson, Michael G. (2011). Palynology. Plant Systematics. Academic Press. pp. 453464. ISBN 978-008-051404-8. Retrieved 6 January 2014.

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 [10] Singh, Gurcharan (2004-01-01). Palynology. Plant
Systematics: An Integrated Approach. p. 142. ISBN
species.[31] Pollen evidence can also reveal the season
[32]
9781578083510.
Retrieved 23 January 2014. In Singh
in which a particular object picked up the pollen.
(2004).
Pollen has been used to trace activity at mass graves
in Bosnia,[33] catch a burglar who brushed against a [11] Judd, Walter S. & Olmstead, Richard G. (2004). A
Hypericum bush during a crime,[34] and has even been
survey of tricolpate (eudicot) phylogenetic relationships.
proposed as an additive for bullets to enable tracking
American Journal of Botany. 91 (10): 16271644.
doi:10.3732/ajb.91.10.1627. PMID 21652313.
them.[35]

[12] Oder, Tom. Dear allergy suerers: Don't blame goldenrod. mnn.com. Mother Nature Network. Retrieved 18
July 2016.
[13] Skoner, DP (July 2001). Allergic rhinitis: denition,
epidemiology, pathophysiology, detection, and diagnosis.. The Journal of allergy and clinical immunology. 108
(1 Suppl): S28. PMID 11449200.
[14] Allergies and Hay Fever WebMD. Retrieved on 2010-0309
[15] Siewertsen, Bjarne. "Hrd nyser for allergikere i varm
fremtid" (English: Hard sneeze for allergic people in
warm future) Danish Meteorological Institute, 18 April
2015. Retrieved: 19 April 2015.
[16] Bee, grass pollen allergy symptoms. allergiesandtreatments.com. Retrieved on 2010-03-09
[17] Van Overtvelt L. et al. Immune mechanisms of allergenspecic sublingual immunotherapy. Revue franaise
dallergologie et dimmunologie clinique. 2006; 46: 713
720.
[18] The Pollen Feeders. Relationships of Natural Enemies and Non-Prey Foods. 7. 2009. pp. 8711.
doi:10.1007/978-1-4020-9235-0_6. ISBN 978-1-40209234-3.
[19] Schwarze, Francis W. M. R.; Engels, Julia; Mattheck,
Claus (2000). Fungal Strategies of Wood Decay in Trees.
Springer. p. 61. ISBN 978-3-540-67205-0.
[20] Salcledo, Christian. Evidence of Pollen Digestion at
Nocturnal Aggregations of Heliconius Sara in Costa Rica
(Lepidoptera: Nymphalidae). Trop. Lepid. Res. 20.1
(2010): 3537. Web.

[25] https://www.youtube.com/watch?v=JBP9pw2rNk4
[26] Graystock, Peter; Yates, Kathryn; Evison, Sophie E. F.;
Darvill, Ben; Goulson, Dave; Hughes, William O. H. (July
2013). The Trojan hives: pollinator pathogens, imported
and distributed in bumblebee colonies. Journal of Applied Ecology: n/an/a. doi:10.1111/1365-2664.12134.
[27] Singh, Rajwinder; Levitt, Abby L.; Rajotte, Edwin
G.; Holmes, Edward C.; Ostiguy, Nancy; vanEngelsdorp, Dennis; Lipkin, W. Ian; 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.
[28] Graystock, Peter; Goulson, Dave; Hughes, William O.
H. (5 August 2015). Parasites in bloom: owers aid
dispersal and transmission of pollinator parasites within
and between bee species. Proceedings of the Royal Society B: Biological Sciences. 282 (1813): 20151371.
doi:10.1098/rspb.2015.1371.
[29] Graystock, Peter; Blane, Edward J.; McFrederick,
Quinn S.; Goulson, Dave; Hughes, William O.H.
(October 2015).
Do managed bees drive parasite spread and emergence in wild bees?". International Journal for Parasitology: Parasites and Wildlife.
doi:10.1016/j.ijppaw.2015.10.001.
[30] Graystock, P.; Jones, J.C.; Pamminger, T.; Parkinson,
J.F.; Norman, V.; Blane, E.J.; Rothstein, L.; Wckers, F.; Goulson, D.; Hughes, W.O.H. (May 2016).
Hygienic food to reduce pathogen risk to bumblebees. Journal of Invertebrate Pathology. 136: 6873.
doi:10.1016/j.jip.2016.03.007.

[21] Cardoso MZ, Gilbert LE; Gilbert (June 2013). Pollen


feeding, resource allocation and the evolution of chemical
defence in passion vine butteries. Journal of Evolutionary Biology. 26 (6): 125460. doi:10.1111/jeb.12119.
PMID 23662837.

[31] Bryant, Vaughn M. Forensic Palynology: A New Way to


Catch Crooks. crimeandclues.com. Archived from the
original on 2007-02-03.

[22] Sanford, Malcolm T. Producing Pollen at the Wayback


Machine (archived January 13, 2007), University of
Florida, Institute of Food and Agricultural Sciences; citing P. Witherell, Other Products of the Hive, Chapter
XVIII, The Hive and the Honey Bee, Dadant & Sons, Inc.,
Hamilton, IL, 1975

[33] Wood, Peter (9 September 2004). Pollen helps war crime


forensics. BBC News.

[23] Mao W, Schuler MA, Berenbaum MR; Schuler; Berenbaum (May 2013). Honey constituents up-regulate
detoxication and immunity genes in the western honey
bee Apis mellifera. Proceedings of the National Academy
of Sciences of the United States of America. 110
(22): 88426. doi:10.1073/pnas.1303884110. PMC
3670375 . PMID 23630255.
[24] Sanford, Malcolm T. Producing Pollen. University
of Florida, Institute of Food and Agricultural Sciences.
Archived from the original on 2007-01-13. Retrieved
2007-08-30.. Document ENY118. Original publication
date November 1, 1994. Revised February 1, 1995. Reviewed May 1, 2003.

[32] Stackhouse, Robert (17 April 2003). Forensics studies


look to pollen. The Battalion.

[34] D. Mildenhall (2006). Hypericum pollen determines the


presence of burglars at the scene of a crime: An example
of forensic palynology. Forensic Science International.
163 (3): 231235. doi:10.1016/j.forsciint.2005.11.028.
PMID 16406430.
[35] Wolf, Lauren K. (18 August 2008). Newscripts. Chemical & Engineering News. 86 (33): 88. doi:10.1021/cenv086n033.p088.

9 Bibliography
Furness, Carol A.; Rudall, Paula J. (January
2001), Pollen and anther characters in monocot systematics, Grana, 40 (12): 1725,
doi:10.1080/00173130152591840

10
Furness, Carol A.; Rudall, Paula J. (November
2003). Apertures with Lids: Distribution and Signicance of Operculate Pollen in Monocotyledons.
International Journal of Plant Sciences. 164 (6):
835854. doi:10.1086/378656.
Davis, Owen (1999). Palynology Pollen. University of Arizona. Department of Geosciences.
Simpson, Michael G. (2011). Plant Systematics.
Academic Press. ISBN 0-08-051404-9. Retrieved
12 February 2014.
Singh, Gurcharan (2004). Plant Systematics: An Integrated Approach. Science Publishers. ISBN 157808-351-6. Retrieved 23 January 2014.

10

External links

Pollen and Spore Identication Literature


Pollen micrographs at SEM and confocal microscope
The ight of a pollen cloud
PalDat (database comprising palynological data
from a variety of plant families)
YouTube video of pollen clouds from Juncus gerardii plants
This article incorporates text from a publication now
in the public domain: Chisholm, Hugh, ed. (1911).
"article name needed ". Encyclopdia Britannica (11th ed.).
Cambridge University Press.

EXTERNAL LINKS

11
11.1

Text and image sources, contributors, and licenses


Text

Pollen Source: https://en.wikipedia.org/wiki/Pollen?oldid=740448875 Contributors: AxelBoldt, Magnus Manske, Mav, Bryan Derksen,
Tarquin, Heron, Jtdirl, Menchi, Arpingstone, Egil, Ellywa, Stan Shebs, Tristanb, KayEss, Marshman, Chrisjj, Pollinator, Robbot, Mayooranathan, Modeha, Dina, Radagast, Giftlite, Graeme Bartlett, Christopher Parham, MPF, Solipsist, Ryanaxp, Utcursch, Dullhunk,
HorsePunchKid, Kate, Solitude, Rich Farmbrough, Bender235, Syp, Elwe, Summer Song, Circeus, Fir0002, Kappa, Nk, Tgabor, PrimEviL,
Danski14, Arthena, Wtmitchell, Dschwen, Velella, Shoey, Gene Nygaard, Hoziron, Richard Arthur Norton (1958- ), Jacobolus, Bratsche,
Polyparadigm, WadeSimMiser, SCEhardt, Allen3, Kesla, Enzo Aquarius, Sjakkalle, Rjwilmsi, Astronaut, Sango123, FlaBot, Ground
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RobotE, Neitherday, Hede2000, Chris Capoccia, Wimt, Tavilis, Anomalocaris, Curtis Clark, Trovatore, VinceBowdren, E rulez, Davemck, Dbrs, CLW, Square87~enwiki, PTSE, KGasso, Esprit15d, Allens, Kungfuadam, SmackBot, Brya, Unyoyega, EncycloPetey, Hardyplants, HalfShadow, Gilliam, Ohnoitsjamie, EdytaT, Chaojoker, Rkitko, SchftyThree, Octahedron80, Nbarth, Mona, A. B., Rlevse, Can't
sleep, clown will eat me, Kingdon, Dreadstar, Richard001, Spentangeli, Thistheman, GameKeeper, Clicketyclack, Anlace, Scientizzle,
Kipala, Mgiganteus1, Ph89~enwiki, Railsmart, Soulkeeper, Smith609, Caiaa, Phuzion, Iridescent, TwistOfCain, Paul venter, Courcelles,
Tawkerbot2, J Milburn, JForget, CmdrObot, Sashag, Corpx, Tawkerbot4, Alexsamson, Omicronpersei8, Dyanega, Rosser1954, PKT,
Sagaciousuk, Missvain, Escarbot, Seaphoto, KP Botany, TimVickers, Smartse, JAnDbot, BlindEagle, Acroterion, Wasell, Magioladitis,
Connormah, VoABot II, Hasek is the best, Mortrek, Michael Goodyear, Catgut, JJ Harrison, Hamiltonstone, Edgecution, WLU, Peter coxhead, Gidip, J.delanoy, Rgoodermote, Thaurisil, NerdyNSK, Vcbio, Icseaturtles, Acalamari, Medium69, Katalaveno, Ratchild, Chiswick
Chap, Richard D. LeCour, NewEnglandYankee, Student7, Nadiatalent, DorganBot, G. Vlcker, Mr Neat, Vlmastra, Philip Trueman,
TXiKiBoT, Oshwah, Mercurywoodrose, Nicholasnice, Malinaccier, A4bot, Someguy1221, Muhammad Mahdi Karim, Raymondwinn,
BotKung, Synthebot, Bonhomme.vincent, Temporaluser, SieBot, ScAvenger lv, JSpung, Oxymoron83, Faradayplank, Steven Crossin,
Evangstadt04, Sunrise, OKBot, Hamiltondaniel, Ppp2k1, Bpeps, Denisarona, ClueBot, CyrilThePig4, Arakunem, Uncle Milty, CounterVandalismBot, Cchba04, Jagdfeld, Alexbot, Jusdafax, SpikeToronto, UrsoBR, SoxBot III, Vanished User 1004, XLinkBot, SwirlBoy39,
Avoided, WikHead, Addbot, DOI bot, Cst17, Bassbonerocks, Numbo3-bot, Wolfeye90, Tide rolls, Lightbot, Cesiumfrog, Luckas-bot,
Yobot, AnakngAraw, AnomieBOT, EryZ, Materialscientist, E235, Citation bot, Wikepedia1515, ArthurBot, Rynejacob, Xqbot, Yazee,
Deanaschwarz, The sock that should not be, Volkan Y, N419BH, FrescoBot, G.Voelcker, Citation bot 1, Pinethicket, Mjs1991, TobeBot, Trappist the monk, Lb.at.wiki, David Hedlund, Arad3333, ChanDMan2010, Aircorn, TGCP, EmausBot, WikitanvirBot, Immunize,
Jadeslair, Wikipelli, Dcirovic, K6ka, Burstwes38, Wikitoov, F, Josve05a, Traxs7, Ovidiucb, Gz33, Wayne Slam, Pun, ChuispastonBot, Forever Dusk, JonRicheld, Biosicherheit, Kleopatra, ClueBot NG, Joebob23456789, A520, PaleCloudedWhite, Widr, Helpful Pixie Bot, Calabe1992, BG19bot, J991, Pbadhikari, Pocketthis, Bombusp, Hamish59, , Persianpollen, 512bits, JYBot, Dexbot,
Shuhrataxmedov, Sminthopsis84, Cerabot~enwiki, Lugia2453, Frosty, Zorahia, JamesMoose, Morganclem, JaconaFrere, Zanbooor, EddBarrows, SirMinkMay, Monkbot, Vieque, Mrpiprabbit, Kinetic37, Davisonio, Arjunnaha, KasparBot, Emely309, Quackriot, Alch348,
Njr24, Trustworthybastile, Fox829, Gods Godzilla and Anonymous: 308

11.2

Images

File:Ambox_globe_content.svg Source: https://upload.wikimedia.org/wikipedia/commons/b/bd/Ambox_globe_content.svg License:


Public domain Contributors: Own work, using File:Information icon3.svg and File:Earth clip art.svg Original artist: penubag
File:Ambox_important.svg Source: https://upload.wikimedia.org/wikipedia/commons/b/b4/Ambox_important.svg License: Public domain Contributors: Own work, based o of Image:Ambox scales.svg Original artist: Dsmurat (talk contribs)
File:Apis_mellifera_flying.jpg Source: https://upload.wikimedia.org/wikipedia/commons/9/99/Apis_mellifera_flying.jpg License:
GFDL 1.2 Contributors: Own work Original artist: Muhammad Mahdi Karim (www.micro2macro.net) Facebook Youtube
File:Arabis_voch1-4.jpg Source: https://upload.wikimedia.org/wikipedia/commons/c/ca/Arabis_voch1-4.jpg License: CC BY-SA 3.0
Contributors: Own work Original artist: Marie Majaura
File:Cactus_flower_pollen.jpg Source: https://upload.wikimedia.org/wikipedia/commons/0/04/Cactus_flower_pollen.jpg License:
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File:Coenocytic_Tetrad.gif Source: https://upload.wikimedia.org/wikipedia/commons/f/f4/Coenocytic_Tetrad.gif License: CC BY-SA
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File:Episyrphus_balteatus_-_head_close-up_(aka).jpg Source: https://upload.wikimedia.org/wikipedia/commons/1/1b/Episyrphus_
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File:Misc_pollen_colorized.jpg Source: https://upload.wikimedia.org/wikipedia/commons/2/2a/Misc_pollen_colorized.jpg License:
Public domain Contributors: Source and public domain notice at Dartmouth Electron Microscope Facility ([1], [2]) Original artist: Dartmouth Electron Microscope Facility, Dartmouth College

10

11

TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

File:Oenothera_speciosa_pollen_200x.jpg Source:
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pollen_200x.jpg License: CC BY-SA 3.0 Contributors: Own work Original artist: ZooFari
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Original artist: ?
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Contributors:
Created from scratch in Adobe Illustrator. Based on Image:Question book.png created by User:Equazcion Original artist:
Tkgd2007
File:RedbudPollen.TIF Source: https://upload.wikimedia.org/wikipedia/commons/c/c1/RedbudPollen.TIF License: CC BY-SA 3.0 Contributors: Own work Original artist: Kleopatra
File:Tulip_Stamen_Tip.jpg Source: https://upload.wikimedia.org/wikipedia/commons/e/ee/Tulip_Stamen_Tip.jpg License: CC BY-SA
3.0 Contributors: Own work Original artist: JJ Harrison (jjharrison89@facebook.com)
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100px-Wiki_letter_w.svg.png 2x' data-le-width='44' data-le-height='44' /></a>
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File:__13.jpg Source: https://upload.wikimedia.org/wikipedia/commons/f/f3/%D0%96%D0%B8%
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artist: Shuhrataxmedov

11.3

Content license

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