Pheromone
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[[Image:Nasinov 9024.JPG|right|thumb|Fanning [[honeybee]] exposes [[Nasonov]] gland (white-at tip of abdomen) releasing pheromone to entice swarm into an empty hive]]
A '''pheromone''' (from [[Greek language|Greek]] ''φέρω'' phero "to bear" + ''‘ορμόνη'' "[[hormone]]") is a [[chemical]] that triggers a natural behavioral response in another member of the same species. There are ''alarm pheromones'', ''food trail pheromones'', ''sex pheromones'', and many others that affect behavior or physiology. Their use among [[insect]]s has been particularly well documented, although many [[vertebrate]]s and [[plant]]s also communicate using pheromones.
== Background ==
The term "pheromone" was introduced by Peter Karlson and Martin Lüscher in 1959, based on the Greek ''pherein '' (to transport) and ''hormone'' (to stimulate). They proposed the term to describe chemical signals from [[conspecific]]s which elicit innate behaviours soon after Butenandt characterized the first such chemical, [[Bombykol]] (a chemically well-characterized pheromone released by the female [[silkworm]] to attract mates).<ref>Karlson, P., Lüscher, M. (1959). Pheromones: a new term for a class of biologically active substances. ''[[Nature]]'' '''183''', 55-56.</ref>
==Types==
;Aggregation pheromones
Produced by one or the other sex, aggregation pheromones attract individuals of both sexes.
;Alarm pheromones
Some species release a volatile substance when attacked by a predator that can trigger flight (in [[aphid]]s) or aggression (in bees) in members of the same species. Pheromones also exist in plants: certain plants emit alarm pheromones when grazed upon, resulting in [[tannin]] production in neighboring plants. These tannins make the plants less appetizing for the [[herbivore]].<ref>J.du P. Bothma, Game ranch management, fourth edition, Van Schaik publishers, 2002</ref>
;Epideictic pheromones
Recognized in insects, epideictic pheromones are different from territory pheromones. According to [[Jean Henri Fabre|Fabre]] (translated from French), "Females who lay their eggs in these fruits deposit these mysterious substances in the vicinity of their clutch to signal to other females of the same species so that they will clutch elsewhere."
[[Image:Bug aggregation.jpg|right|thumb|Aggregation of bug nymphs]]
;Releaser pheromones
Releaser pheromones are powerful attractant molecules that some organisms may use to attract mates from a distance of two miles or more. This type of pheromone generally elicits a rapid response but is quickly degraded. In contrast, a primer pheromone has a slower onset and a longer duration.
;Primer pheromones
Primer pheromones trigger a change of developmental events.
;Territorial pheromones
Laid down in the environment, territorial pheromones mark the boundaries of an organism's territory. In dogs, these hormones are present in the urine, which they deposit on landmarks serving to mark the perimeter of the claimed territory.
;Trail pheromones
Trail pheromones are common in social insects. For example, [[ant]]s mark their paths with these pheromones, which are non-volatile [[hydrocarbon]]s.
Certain ants lay down an initial trail of pheromones as they return to the nest with food. This trail attracts other ants and serves as a guide.<ref>{{cite web |url=http://www.news.cornell.edu/releases/Feb98/antpheromone.hrs.html | title=Excited ants follow pheromone trail of same chemical they will use to paralyze their prey | accessdate=2006-03-14}}</ref> As long as the food source remains, the pheromone trail will be continually renewed. The pheromone must be continually renewed because it evaporates quickly. When the supply begins to dwindle, the trailmaking ceases. In at least one species of ant, trails that no longer lead to food are also marked with a repellent pheromone.<ref>{{cite web | url=http://animal.discovery.com/news/afp/20051128/ants.html | title=Study: Ants Use Scents Like Road Signs | accessdate=2006-03-14}}</ref>
;Sex pheromones
[[Image:Sesiidae Pheromon fg01.jpg|thumb|250px|right|[[Sesiidae]] on a pheromone trap]]
In animals, sex pheromones indicate the availability of the female for breeding. Male animals may also emit pheromones that convey information about their species and [[genotype]]. Many insect species release sex pheromones to attract a mate, and many [[lepidoptera]]ns can detect a potential mate from as far away as 10 kilometers (6 mi). Pheromones can be used in [[gamete]]s to trail the opposite sex's gametes for [[fertilization]]. Pheromones are also used in the detection of [[oestrus]] in [[Pig|sows]]. [[Boar]] pheromones are sprayed into the [[wiktionary:sty|sty]], and those sows which exhibit [[sexual arousal]] are known to be currently available for breeding.
;Other pheromones (not yet classified)
This classification, based on the effects on behavior, remains artificial. Pheromones fill many additional functions.
* [[Nasonov]] pheromones (worker bees)
* Royal pheromones (bees)
* Calming (appeasement) pheromones (mammals)
==Uses==
===Animals===
Pheromones of the pest insect species, such as the [[Japanese beetle]] and the [[gypsy moth]], can be used to induce many behaviors. This facilitates trapping for monitoring purposes and population control by creating confusion, disrupting mating and preventing them from laying eggs.
In [[mammal]]s and [[reptile]]s, pheromones may be detected by the [[vomeronasal organ]], or Jacobson's organ, which lies between the nose and mouth and is the first stage of the [[accessory olfactory system]]. Some pheromones in these animals are detected by regular [[olfactory]] membranes.
===Humans===
Few well-controlled scientific studies have ever been published suggesting the possibility of pheromones in humans. The best known case involves the reported synchronization of [[menstrual cycle]]s among women based on unconscious odor cues (the ''[[McClintock effect]]'', named after the primary investigator, [[Martha McClintock]], of the University of Chicago).<ref>McClintock MK (1971). "Menstrual synchrony and suppression". Nature 229 (5282): 244-5. PMID 4994256</ref><ref>Stern K, McClintock MK (1998). "Regulation of ovulation by human pheromones". Nature 392 (6672): 177-9. doi:10.1038/32408. PMID 9515961.</ref> This study proposes that there are two types of pheromone involved: "One, produced prior to ovulation, shortens the ovarian cycle; and the second, produced just at ovulation, lengthens the cycle". However recent studies and reviews of the of the McClintock methodology have called into question the validity of her results. <ref>{{cite journal | last = Yang | first = Zhengwei | coauthors = Jeffrey C. Schank | title = Women Do Not Synchronize Their Menstrual Cycles | journal = Human Nature | volume = 17 | issue = 4 | pages = 434–447 | year = 2006 | url = http://transactionpub.metapress.com/openurl.asp?genre=article&issn=1045-6767&volume=17&issue=4&spage=434 | accessdate = 2007-06-25 | doi = 10.1007/s12110-006-1005-z }}</ref>
Other studies have suggested that people might be using odor cues associated with the immune system to [[Disassortative sexual selection|select mates who are not closely related to themselves]]. Using a brain imaging technique, [[Sweden|Swedish]] researchers have shown that [[Homosexuality|homosexual]] and [[Heterosexuality|heterosexual]] males' brains respond differently to two odors that may be involved in sexual arousal, and that the homosexual men respond in the same way as heterosexual women. The study was expanded to include homosexual women; the results were consistent with previous findings meaning that homosexual women were not as responsive to male identified odors, but their response to female cues was similar to heterosexual males.<ref> Savic, I."[http://www.pnas.org/cgi/content/abstract/0600331103v1?maxtoshow=&HITS=10&hits=10&RESULTFORMAT=&fulltext=ivanka&searchid=1&FIRSTINDEX=0&resourcetype=HWCIT Brain response to putative pheromones in lesbian women]." PNAS, May 16, 2006</ref> According to the researchers, this research suggests a possible role for human pheromones in the biological basis of [[sexual orientation]].<ref>Wade, N. "[http://www.nytimes.com/2005/05/09/science/09cnd-smell.html Gay Men are found to have Different Scent of Attraction]." NY Times, May 9, 2005</ref>
Another study demonstrated that the smell of [[androstadienone]], a chemical component of male [[sweat]], maintains higher levels of [[cortisol]] in females. The scientists suggest that the ability of this compound to influence the [[endocrine]] balance of the opposite sex makes it a human pheromonal chemosignal.<ref>Wyart C, Webster WW, Chen JH, Wilson SR, McClary A, Khan RM, Sobel N. 2007. Smelling a single component of male sweat alters levels of cortisol in women. J Neurosci. 27(6):1261-5.[http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=pubmed&cmd=Retrieve&dopt=AbstractPlus&list_uids=17287500&query_hl=1&itool=pubmed_docsum]</ref> In 2002 a study published in the quarterly journal ''Physiology and Behavior'' showed an unnamed synthetic chemical in women's perfume appeared to increase intimate contact with men. The authors hypothesize, but do not demonstrate, that the observed behavioural differences are olfactory mediated. <ref name = "sf"> {{cite journal
| first =
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| year =2002
| month =March 20
| title =San Francisco State University study shows that synthetic pheromones in women's perfume increase intimate contact with men
| journal =San Francisco State University Office of Public Affairs
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| url =http://www.sfsu.edu/~news/prsrelea/fy01/091.htm
}} </ref>
In 2006 it was shown that a second mouse [[odorant receptor|receptor]] sub-class is found in the [[olfactory epithelium]]. Called the [[trace amine-associated receptor]]s (TAAR), some are activated by volatile [[amine]]s found in mouse urine, including one putative mouse pheromone.<ref>Liberles SD, Buck LB. 2006. A second class of chemosensory receptors in the olfactory epithelium. Nature. 442(7103):645-50. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=pubmed&cmd=Retrieve&dopt=AbstractPlus&list_uids=16878137&query_hl=1&itool=pubmed_docsum]</ref> [[Ortholog]]ous receptors exist in humans providing, the authors propose, evidence for a mechanism of human pheromone detection.<ref>Pearson H. 2006. Mouse data hint at human pheromones. Nature. 442(7102):495. [http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=pubmed&cmd=Retrieve&dopt=AbstractPlus&list_uids=16885951&query_hl=5&itool=pubmed_docsum]</ref>
Some body spray advertisers claim that their products contain human sexual pheromones which act as an [[aphrodisiac]]. In the 1970's "copulins" were patented as products which release human pheromones, based on research on [[rhesus monkey]]s.<ref name="wy">Wyatt, Tristram D. (2003). ''Pheromones and Animal Behaviour: Communication by Smell and Taste.'' Cambridge: Cambridge University Press. ISBN 0-521-48526-6. p. 298 Quoting Preti & Weski (1999) "No peer reviewed data supporting the presences of...human...pheromones that cause rapid behavioral changes, such as attraction and/or [[copulation]] have been documented." </ref> Subsequently, androstenone, axillary sweat, and "vomodors" have been claimed to act as human pheromones.<ref name = "Warren S. T. Hays">Hays, Warren S. T., Human pheromones: have they been demonstrated? ''Behavioral Ecology and Sociobiology'', 2003, 54:89-97</ref> Despite these claims, no pheromonal substance has ever been demonstrated to directly influence human behavior in a [[peer review]]ed study.<ref name="wy" /><ref name = "Neu"> {{cite book
| last =Bear
| first =Mark F.
| authorlink =
| coauthors =Barry W. Connors, Michael A. Paradiso
| year =2006
| title =Neuroscience: Exploring the Brain
| publisher =
| location =
| id =ISBN 0781760038
}} p. 264 ...there has not yet been any hard evidence for human pheromones that might [change] sexual attraction (for members of either sex) [naturally] </ref><ref name = "Warren S. T. Hays"/>
==See also==
<div style="-moz-column-count:2; column-count:2;">
* [[Allomone]]
* [[Androstenol]]
* [[Androstenone]]
* [[Body odor]]
* [[Claus Wedekind]]
* [[Estratetraenol]]
* [[Feline facial pheromone]]
* [[Honey bee pheromones]]
* [[Kairomone]]
* [[Mating Disruption]]
* [[Osmeterium]], an organ in swallowtail caterpillars
* [[Pheromones in fiction]]
* [[Quorum sensing]]
* [[Semiochemical]]
* [[Sweat]]
* [[Testosterone]]
* [[Vomeronasal organ]]
</div>
==References==
{{reflist}}
==Further reading==
*Kohl, JV., Atzmueller, M., Fink, B. & Grammer, K. (2001). Human Pheromones: Integrating Neuroendocrinology and Ethology. ''Neuroendocrinology Letters, 22(5),'' 319-331. [http://evolution.anthro.univie.ac.at/institutes/urbanethology/resources/pdf/NEL220501R01_.pdf Full text]
*Liberles, S.D., [[Linda Buck|Buck, L.B.]] (2006). A second class of chemosensory receptors in the olfactory epithelium. ''Nature, 442,'' 645-50.
*McClintock, M.K. (1984). Estrous synchrony: modulation of ovarian cycle length by female pheromones. ''Physiological Behavior, 32,'' 701-705.
*[[Edward O. Wilson|Wilson, E. O.]], Bossert, W. H. (1963). Chemical communication among animals. ''Recent Progress in Hormone Research, 19,'' 673-716.
*Wyatt, Tristram D. (2003). ''Pheromones and Animal Behaviour: Communication by Smell and Taste.'' Cambridge: Cambridge University Press. ISBN 0-521-48526-6.
==External links==
*[http://www.pherobase.com/ Pherobase], the database of insect pheromones
*[http://www.biolreprod.org/cgi/content/full/68/6/2107 Male Axillary Extracts Contain Pheromones that Affect Pulsatile Secretion of Luteinizing Hormone and Mood in Women Recipients]
*[http://www.cbsnews.com/stories/2005/05/09/tech/main694078.shtml Sexual Orientation, in the Brain]
*[http://www.ondwelle.com/OSM03.pdf Review of published debate about HOW insects detect pheromones over large distances, even when the wind seems unfavourable]
*[http://www.berkeley.edu/news/media/releases/2007/02/06_sweat.shtml Male sweat boosts women's hormone levels] -- from [[UC Berkeley]], February 2007
*[http://www.sciencedaily.com/releases/2007/02/070207172019.htm The Effect of Male Sweat on Women's Hormone Levels] -- from ''Science Daily'', February 2007
*[http://www.sciencedaily.com/releases/2003/03/030317074228.htm Pheromones In Male Perspiration Reduce Women's Tension, Alter Hormone Response] -- from ''Science Daily'' (March 2003)
[[Category:Pheromones|*]]
[[Category:Greek loanwords]]
[[ar:فيرومون]]
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