A pheromone (from Ancient Greek φέρω (phérō) 'to bear', and hormone) is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavior of the receiving individuals.[1] There are alarm pheromones, food trail pheromones, sex pheromones, and many others that affect behavior or physiology. Pheromones are used by many organisms, from basic unicellular prokaryotes to complex multicellular eukaryotes.[2] Their use among insects has been particularly well documented. In addition, some vertebrates, plants and ciliates communicate by using pheromones. The ecological functions and evolution of pheromones are a major topic of research in the field of chemical ecology.[3]

A fanning honeybee exposes Nasonov's gland (white – at tip of abdomen) releasing pheromone to entice swarm into an empty hive



The portmanteau word "pheromone" was coined by Peter Karlson and Martin Lüscher in 1959, based on the Greek φέρω phérō ('I carry') and ὁρμων hórmōn ('stimulating').[4] Pheromones are also sometimes classified as ecto-hormones. They were researched earlier by various scientists, including Jean-Henri Fabre, Joseph A. Lintner, Adolf Butenandt, and ethologist Karl von Frisch who called them various names, like for instance "alarm substances". These chemical messengers are transported outside of the body and affect neurocircuits, including the autonomous nervous system with hormone or cytokine mediated physiological changes, inflammatory signaling, immune system changes and/or behavioral change in the recipient.[5] They proposed the term to describe chemical signals from conspecifics that elicit innate behaviors soon after the German biochemist Adolf Butenandt had characterized the first such chemical, bombykol, a chemically well-characterized pheromone released by the female silkworm to attract mates.[6]

Categorization by function



Aggregation of bug nymphs
Aggregation of the water springtail Podura aquatica

Aggregation pheromones function in mate choice, overcoming host resistance by mass attack, and defense against predators. A group of individuals at one location is referred to as an aggregation, whether consisting of one sex or both sexes. Male-produced sex attractants have been called aggregation pheromones, because they usually result in the arrival of both sexes at a calling site and increase the density of conspecifics surrounding the pheromone source. Most sex pheromones are produced by the females; only a small percentage of sex attractants are produced by males.[7] Aggregation pheromones have been found in members of the Coleoptera, Collembola,[8] Diptera, Hemiptera, Dictyoptera, and Orthoptera. In recent decades, aggregation pheromones have proven useful in the management of many pests, such as the boll weevil (Anthonomus grandis), the pea and bean weevil (Sitona lineatus, and stored product weevils (e.g. Sitophilus zeamais, Sitophilus granarius, and Sitophilus oryzae). Aggregation pheromones are among the most ecologically selective pest suppression methods. They are non-toxic and effective at very low concentrations.[9]



Some species release a volatile substance when attacked by a predator that can trigger flight (in aphids) or aggression (in ants, bees, termites, and wasps)[10][11][12][13][14] in members of the same species. For example, Vespula squamosa use alarm pheromones to alert others to a threat.[15] In Polistes exclamans, alarm pheromones are also used as an alert to incoming predators.[16] Pheromones also exist in plants: Certain plants emit alarm pheromones when grazed upon, resulting in tannin production in neighboring plants.[17] These tannins make the plants less appetizing to herbivores.[17]

An alarm pheromone has been documented in a mammalian species. Alarmed pronghorn, Antilocapra americana flair their white rump hair and exposes two highly odoriferous glands that releases a compound described having the odor "reminiscent of buttered popcorn". This sends a message to other pronghorns by both sight and smell about a present danger. This scent has been observed by humans 20 to 30 meters downwind from alarmed animals. The major odour compound identified from this gland is 2-pyrrolidinone.[18]



Epideictic pheromones are different from territory pheromones, when it comes to insects. Fabre observed and noted how "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 they should clutch elsewhere." It may be helpful to note that the word epideictic, having to do with display or show (from the Greek 'deixis'), has a different but related meaning in rhetoric, the human art of persuasion by means of words.



Laid down in the environment, territorial pheromones mark the boundaries and identity of an organism's territory. Cats and dogs deposit these pheromones by urinating on landmarks that mark the perimeter of the claimed territory. In social seabirds, the preen gland is used to mark nests, nuptial gifts, and territory boundaries with behavior formerly described as 'displacement activity'.[19]



Social insects commonly use trail pheromones. For example, ants mark their paths with pheromones consisting of volatile hydrocarbons. 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.[20] As long as the food source remains available, visiting ants will continuously renew the pheromone trail. The pheromone requires continuous renewal because it evaporates quickly. When the food supply begins to dwindle, the trail-making ceases. Pharaoh ants (Monomorium pharaonis) mark trails that no longer lead to food with a repellent pheromone, which causes avoidance behaviour in ants.[21] Repellent trail markers may help ants to undertake more efficient collective exploration.[22] The army ant Eciton burchellii provides an example of using pheromones to mark and maintain foraging paths. When species of wasps such as Polybia sericea found new nests, they use pheromones to lead the rest of the colony to the new nesting site.

Gregarious caterpillars, such as the forest tent caterpillar, lay down pheromone trails that are used to achieve group movement.[23]

Male Danaus chrysippus showing the pheromone pouch and brush-like organ in Kerala, India

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.

At the microscopic level, a number of bacterial species (e.g. Bacillus subtilis, Streptococcus pneumoniae, Bacillus cereus) release specific chemicals into the surrounding media to induce the "competent" state in neighboring bacteria.[24] Competence is a physiological state that allows bacterial cells to take up DNA from other cells and incorporate this DNA into their own genome, a sexual process called transformation.

Among eukaryotic microorganisms, pheromones promote sexual interaction in numerous species.[25] These species include the yeast Saccharomyces cerevisiae, the filamentous fungi Neurospora crassa and Mucor mucedo, the water mold Achlya ambisexualis, the aquatic fungus Allomyces macrogynus, the slime mold Dictyostelium discoideum, the ciliate protozoan Blepharisma japonicum and the multicellular green algae Volvox carteri. In addition, male copepods can follow a three-dimensional pheromone trail left by a swimming female, and male gametes of many animals use a pheromone to help find a female gamete for fertilization.[26]

Many well-studied insect species, such as the ant Leptothorax acervorum, the moths Helicoverpa zea and Agrotis ipsilon, the bee Xylocopa sonorina, the frog Pseudophryne bibronii, and the butterfly Edith's checkerspot release sex pheromones to attract a mate, and some lepidopterans (moths and butterflies) can detect a potential mate from as far away as 10 km (6.2 mi).[27][28] Some insects, such as ghost moths, use pheromones during lek mating.[29] Traps containing pheromones are used by farmers to detect and monitor insect populations in orchards. In addition, Colias eurytheme butterflies release pheromones, an olfactory cue important for mate selection.[30] In mealworm beetles, Tenebrio molitor, the female preference of pheromones is dependent on the nutritional condition of the males.

The effect of Hz-2V virus infection on the reproductive physiology and behavior of female Helicoverpa zea moths is that in the absence of males they exhibited calling behavior and called as often but for shorter periods on average than control females. Even after these contacts virus-infected females made many frequent contacts with males and continued to call; they were found to produce five to seven times more pheromone and attracted twice as many males as did control females in flight tunnel experiments.[31]

Pheromones are also utilized by bee and wasp species. Some pheromones can be used to suppress the sexual behavior of other individuals allowing for a reproductive monopoly – the wasp R. marginata uses this.[32] With regard to the Bombus hyperboreus species, males, otherwise known as drones, patrol circuits of scent marks (pheromones) to find queens.[33] In particular, pheromones for the Bombus hyperboreus, include octadecenol, 2,3-dihydro-6-transfarnesol, citronellol, and geranylcitronellol.[34]

Sea urchins release pheromones into the surrounding water, sending a chemical message that triggers other urchins in the colony to eject their sex cells simultaneously.

In plants, some homosporous ferns release a chemical called antheridiogen, which affects sex expression. This is very similar to pheromones.



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)
  • Necromones, given off by a deceased and decomposing organism; consisting of oleic and linoleic acids, they allow crustaceans and hexapods to identify the presence of dead conspecifics.[35]
  • Suckling: TAA is present in rabbit milk and seems to play a role of pheromone inducing suckling in the newborn rabbit.[36]

Categorization by type




Releaser pheromones are pheromones that cause an alteration in the behavior of the recipient. For example, some organisms use powerful attractant molecules to attract mates from a distance of two miles or more. In general, this type of pheromone elicits a rapid response, but is quickly degraded. In contrast, a primer pheromone has a slower onset and a longer duration. For example, rabbit (mothers) release mammary pheromones that trigger immediate nursing behavior by their babies.[19]



Primer pheromones trigger a change of developmental events (in which they differ from all the other pheromones, which trigger a change in behavior). They were first described in Schistocerca gregaria by Maud Norris in 1954.[37]



Signal pheromones cause short-term changes, such as the neurotransmitter release that activates a response. For instance, GnRH molecule functions as a neurotransmitter in rats to elicit lordosis behavior.[5]

Pheromone receptors


In the olfactory epithelium


The human trace amine-associated receptors are a group of six G protein-coupled receptors (i.e., TAAR1, TAAR2, TAAR5, TAAR6, TAAR8, and TAAR9) that – with exception for TAAR1 – are expressed in the human olfactory epithelium.[38] In humans and other animals, TAARs in the olfactory epithelium function as olfactory receptors that detect volatile amine odorants, including certain pheromones;[38][39] these TAARs putatively function as a class of pheromone receptors involved in the olfactive detection of social cues.[38][39]

A review of studies involving non-human animals indicated that TAARs in the olfactory epithelium can mediate attractive or aversive behavioral responses to a receptor agonist.[39] This review also noted that the behavioral response evoked by a TAAR can vary across species (e.g., TAAR5 mediates attraction to trimethylamine in mice and aversion to trimethylamine in rats).[39] In humans, hTAAR5 presumably mediates aversion to trimethylamine, which is known to act as an hTAAR5 agonist and to possess a foul, fishy odor that is aversive to humans;[39][40] however, hTAAR5 is not the only olfactory receptor that is responsible for trimethylamine olfaction in humans.[39][40] As of December 2015, hTAAR5-mediated trimethylamine aversion has not been examined in published research.[40]

In the vomeronasal organ


In reptiles, amphibia and non-primate mammals pheromones are detected by regular olfactory membranes, and also by the vomeronasal organ (VNO), or Jacobson's organ, which lies at the base of the nasal septum between the nose and mouth and is the first stage of the accessory olfactory system.[41] While the VNO is present in most amphibia, reptiles, and non-primate mammals,[42] it is absent in birds, adult catarrhine monkeys (downward facing nostrils, as opposed to sideways), and apes.[43] An active role for the human VNO in the detection of pheromones is disputed; while it is clearly present in the fetus it appears to be atrophied, shrunk or completely absent in adults. Three distinct families of vomeronasal receptors, putatively pheromone sensing, have been identified in the vomeronasal organ named V1Rs, V2Rs, and V3Rs. All are G protein-coupled receptors but are only distantly related to the receptors of the main olfactory system, highlighting their different role.[41]



Olfactory processing of chemical signals like pheromones exists in all animal phyla and is thus the oldest of the senses.[citation needed] It has been suggested that it serves survival by generating appropriate behavioral responses to the signals of threat, sex and dominance status among members of the same species.[44]

Furthermore, it has been suggested that in the evolution of unicellular prokaryotes to multicellular eukaryotes, primordial pheromone signaling between individuals may have evolved to paracrine and endocrine signaling within individual organisms.[45]

Some authors assume that approach-avoidance reactions in animals, elicited by chemical cues, form the phylogenetic basis for the experience of emotions in humans.[46]

Evolution of sex pheromones


Avoidance of inbreeding


Mice can distinguish close relatives from more distantly related individuals on the basis of scent signals,[47] which enables them to avoid mating with close relatives and minimizes deleterious inbreeding.[48]

In addition to mice, two species of bumblebee, in particular Bombus bifarius and Bombus frigidus, have been observed to use pheromones as a means of kin recognition to avoid inbreeding.[49] For example, B. bifarius males display "patrolling" behavior in which they mark specific paths outside their nests with pheromones and subsequently "patrol" these paths.[49] Unrelated reproductive females are attracted to the pheromones deposited by males on these paths, and males that encounter these females while patrolling can mate with them.[49] Other bees of the Bombus species are found to emit pheromones as precopulatory signals, such as Bombus lapidarius.[50]



Pheromone trapping


Pheromones of certain pest insect species, such as the Japanese beetle, acrobat ant, and the spongy moth, can be used to trap the respective insect for monitoring purposes, to control the population by creating confusion, to disrupt mating, and to prevent further egg laying.

Animal husbandry


Pheromones are used in the detection of oestrus in sows. Boar pheromones are sprayed into the sty, and those sows that exhibit sexual arousal are known to be currently available for breeding.

Human sex pheromone controversies


While humans are highly dependent upon visual cues, when in close proximity smells also play a role in sociosexual behaviors. An inherent difficulty in studying human pheromones is the need for cleanliness and odorlessness in human participants.[51] Though various researchers have investigated the possibility of their existence, no pheromonal substance has ever been demonstrated to directly influence human behavior in a peer reviewed study.[52][53][54][55] Experiments have focused on three classes of possible human pheromones: axillary steroids, vaginal aliphatic acids, and stimulators of the vomeronasal organ, including this 2018 study claiming pheromones affect men's sexual cognition.

Axillary steroids


Axillary steroids are produced by the testes, ovaries, apocrine glands, and adrenal glands.[56] These chemicals are not biologically active until puberty when sex steroids influence their activity.[57] The change in activity during puberty suggest that humans may communicate through odors.[56] Several axillary steroids have been described as possible human pheromones: androstadienol, androstadienone, androstenol, androstenone, and androsterone.

  • Androstenol is the putative female pheromone.[57] In a 1978 study by Kirk-Smith, people wearing surgical masks treated with androstenol or untreated were shown pictures of people, animals and buildings and asked to rate the pictures on attractiveness.[58] Individuals with their masks treated with androstenol rated their photographs as being "warmer" and "more friendly".[58] The best-known case study involves the synchronization of menstrual cycles among women based on unconscious odor cues, the McClintock effect, named after the primary investigator, Martha McClintock, of the University of Chicago.[59][60] A group of women were exposed to a whiff of perspiration from other women. Depending on the time in the month the sweat was collected (before, during, or after ovulation) there was an association with the recipient woman's menstrual cycle to speed up or slow down. The 1971 study proposed 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 methodology have called the validity of her results and existence of menstrual synchronization into question.[61][62]
  • Androstenone is postulated to be secreted only by males as an attractant for women, and thought to be a positive effector for their mood. It seems to have different effects on women, depending on where a female is in her menstrual cycle, with the highest sensitivity to it during ovulation.[57] In 1983, study participants exposed to androstenone were shown to undergo changes in skin conductance.[63] Androstenone has been found to be perceived as more pleasant to women during their time of ovulation.[51]
  • Androstadienone seems to affect the limbic system and causes a positive reaction in women, improving mood.[56] Responses to androstadienone depend on the individual and the environment they are in.[64] Androstadienone negatively influences[how?] the perception of pain in women.[64] Women tend to react positively after androstadienone presentation, while men react more negatively. In an experiment by Hummer and McClintock, androstadienone or a control odor was put on the upper lips of fifty males and females and they were tested for four effects of the pheromone: 1) automatic attention towards positive and negative facial expressions, 2) the strength of cognitive and emotional information as distractors in a simple reaction time task, 3) relative attention to social and nonsocial stimuli (i.e. neutral faces), and 4) mood and attentiveness in the absence of social interaction. Those treated with androstadienone drew more attention to towards emotional facial expressions and emotional words but no increased attention to neutral faces. These data suggest that androstadienone may increase attention to emotional information causing the individual to feel more focused. It is thought that androstadienone modulates on how the mind attends and processes information.[64]

While it may be expected on evolutionary grounds that humans have pheromones, these three molecules have yet to be rigorously proven to act as such. Research in this field has suffered from small sample sizes, publication bias, false positives, and poor methodology.[65]

Vaginal aliphatic acids


A class of aliphatic acids (volatile fatty acids as a kind of carboxylic acid) was found in female rhesus monkeys that produced six types in the vaginal fluids.[66] The combination of these acids is referred to as "copulins". One of the acids, acetic acid, was found in all of the sampled female's vaginal fluid.[66] Even in humans, one-third of women have all six types of copulins, which increase in quantity before ovulation.[66] Copulins are used to signal ovulation; however, as human ovulation is concealed it is thought that they may be used for reasons other than sexual communication.[56]

Stimulators of the vomeronasal organ


The human vomeronasal organ has epithelia that may be able to serve as a chemical sensory organ; however, the genes that encode the VNO receptors are nonfunctional pseudogenes in humans.[51] Also, while there are sensory neurons in the human VNO there seem to be no connections between the VNO and the central nervous system. The associated olfactory bulb is present in the fetus, but regresses and vanishes in the adult brain. There have been some reports that the human VNO does function, but only responds to hormones in a "sex-specific manner". There also have been pheromone receptor genes found in olfactory mucosa.[51] There have been no experiments that compare people lacking the VNO, and people that have it. It is disputed on whether the chemicals are reaching the brain through the VNO or other tissues.[56]

In 2006, it was shown that a second mouse receptor sub-class is found in the olfactory epithelium. Called the trace amine-associated receptors (TAAR), some are activated by volatile amines found in mouse urine, including one putative mouse pheromone.[67] Orthologous receptors exist in humans providing, the authors propose, evidence for a mechanism of human pheromone detection.[68]

Although there are disputes about the mechanisms by which pheromones function, there is evidence that pheromones do affect humans.[69] Despite this evidence, it has not been conclusively shown that humans have functional pheromones. Those experiments suggesting that certain pheromones have a positive effect on humans are countered by others indicating they have no effect whatsoever.[56]

A possible theory being studied now is that these axillary odors are being used to provide information about the immune system. Milinski and colleagues found that the artificial odors that people chose are determined in part by their major histocompatibility complexes (MHC) combination.[70] Information about an individual's immune system could be used as a way of "sexual selection" so that the female could obtain good genes for her offspring.[51] Claus Wedekind and colleagues found that both men and women prefer the axillary odors of people whose MHC is different from their own.[71]

Some body spray advertisers claim that their products contain human sexual pheromones that act as an aphrodisiac. Despite these claims, no pheromonal substance has ever been demonstrated to directly influence human behavior in a peer reviewed study.[56][54][disputeddiscuss] Thus, the role of pheromones in human behavior remains speculative and controversial.[72]

See also



  1. ^ "Definition of pheromone". Medicinenet. MedicineNet Inc. 19 March 2012. Archived from the original on 11 May 2011. Retrieved 14 February 2010.
  2. ^ Kleerebezem M, Quadri LE (October 2001). "Peptide pheromone-dependent regulation of antimicrobial peptide production in Gram-positive bacteria: a case of multicellular behavior". Peptides. 22 (10): 1579–1596. doi:10.1016/S0196-9781(01)00493-4. PMID 11587786. S2CID 38943224.
  3. ^ Wood William F. (1983). "Chemical Ecology: Chemical Communication in Nature". Journal of Chemical Education. 60 (7): 1531–539. Bibcode:1983JChEd..60..531W. doi:10.1021/ed060p531.
  4. ^ Karlson P, Luscher M (January 1959). "Pheromones': a new term for a class of biologically active substances". Nature. 183 (4653): 55–56. Bibcode:1959Natur.183...55K. doi:10.1038/183055a0. PMID 13622694. S2CID 4243699.
  5. ^ a b Kohl JV, Atzmueller M, Fink B, Grammer K (October 2001). "Human pheromones: integrating neuroendocrinology and ethology". Neuro Endocrinology Letters. 22 (5): 309–321. PMID 11600881.
  6. ^ Butenandt A, Beckmann R, Hecker E (May 1961). "[On the sexattractant of silk-moths. I. The biological test and the isolation of the pure sex-attractant bombykol]". Hoppe-Seyler's Zeitschrift für Physiologische Chemie. 324: 71–83. doi:10.1515/bchm2.1961.324.1.71. PMID 13689417.
  7. ^ "Insect aggregation pheromones". www.msu.edu. Retrieved 19 February 2018.
  8. ^ Salmon S, Rebuffat S, Prado S, Sablier M, d'Haese C, Sun JS, Ponge JF (2019-05-20). "Chemical communication in springtails: a review of facts and perspectives" (PDF). Biology and Fertility of Soils. 55 (5): 425–438. Bibcode:2019BioFS..55..425S. doi:10.1007/s00374-019-01365-8. ISSN 0178-2762. S2CID 159042283.
  9. ^ Landolt JP (1997). "Sex attractant and aggregation pheromones of male phytophagous insects". American Entomologist. 43 (1): 12–22. doi:10.1093/ae/43.1.12.
  10. ^ Sobotník J, Hanus R, Kalinová B, Piskorski R, Cvacka J, Bourguignon T, Roisin Y (April 2008). "(E,E)-alpha-farnesene, an alarm pheromone of the termite Prorhinotermes canalifrons". Journal of Chemical Ecology. 34 (4): 478–486. Bibcode:2008JCEco..34..478S. CiteSeerX doi:10.1007/s10886-008-9450-2. PMID 18386097. S2CID 8755176.
  11. ^ Wood, William F.; Chong, Berni (1975). "3-Octanone and 3-Octanol; Alarm Pheromones from East African Acacia Ants". Journal of the Georgia Entomological Society. 10: 332–334.
  12. ^ Wood, William F.; Palmer, Todd M.; Stanton, Maureen L. (2002). "A comparison of volatiles in mandibular glands from three Crematogaster ant symbionts of the whistling thorn acacia". J Biochemical Systematics and Ecology. 30 (3): 217–222. Bibcode:2002BioSE..30..217W. doi:10.1016/S0305-1978(01)00099-0.
  13. ^ Wood, William F.; Hoang, Thuy-Tien; McGlynn, Terrence P. (2011). "Volatile components from the mandibular glands of the turtle ants, Cephalotes alfaroi and C. cristatus". Biochemical Systematics and Ecology. 39: 135–138. doi:10.1016/j.bse.2011.01.013.
  14. ^ Wood, William F. (2005). "Comparison of mandibular gland volatiles from ants of the bull horn acacia, Acacia collinsii". Biochemical Systematics and Ecology. 33 (7): 651–658. Bibcode:2005BioSE..33..651W. doi:10.1016/j.bse.2004.12.009.
  15. ^ Landoldt, P. J., Reed, H. C., and Heath, R. R. "An Alarm Pheromone from Heads of Worker Vespula squamosa (Hymenoptera: Vespidae)", "Florida Entomologist", June 1999.
  16. ^ Post DC, Downing HA, Jeanne RL (October 1984). "Alarm response to venom by social waspsPolistes exclamans andP. fuscatus (Hymenoptera: Vespidae)". Journal of Chemical Ecology. 10 (10): 1425–1433. Bibcode:1984JCEco..10.1425P. doi:10.1007/BF00990313. PMID 24318343. S2CID 38398672.
  17. ^ a b Marcus JB (2019). Aging, nutrition and taste nutrition, food science and culinary perspectives for aging tastefully. [Place of publication not identified]: Elsevier Academic Press. ISBN 978-0-12-813528-0. OCLC 1097958893.
  18. ^ Wood, William F. (2002). "2-Pyrrolidinone, a putative alerting pheromone from rump glands of pronghorn, Antilocapra Americana". Biochemical Systematics and Ecology. 30 (4): 361–363. Bibcode:2002BioSE..30..361W. doi:10.1016/S0305-1978(01)00097-7.
  19. ^ a b "Kimball, J.W. Pheromones. Kimball's Biology Pages. Sep 2008". Archived from the original on 2018-01-21. Retrieved 2008-11-01.
  20. ^ "Excited ants follow pheromone trail of same chemical they will use to paralyze their prey". Retrieved 2006-03-14.
  21. ^ Robinson EJ, Green KE, Jenner EA, Holcombe M, Ratnieks FL (2008). "Decay rates of attractive and repellent pheromones in an ant foraging trail network" (PDF). Insectes Sociaux. 55 (3): 246–251. doi:10.1007/s00040-008-0994-5. S2CID 27760894.
  22. ^ Hunt ER, Franks NR, Baddeley RJ (June 2020). "The Bayesian superorganism: externalized memories facilitate distributed sampling". Journal of the Royal Society, Interface. 17 (167): 20190848. doi:10.1098/rsif.2019.0848. PMC 7328406. PMID 32546115.
  23. ^ Fitzgerald TD (July 2008). "Use of pheromone mimic to cause the disintegration and collapse of colonies of tent caterpillars ( Malacosoma spp.)". Journal of Applied Entomology. 132 (6): 451–460. doi:10.1111/j.1439-0418.2008.01286.x. S2CID 83824574.
  24. ^ Bernstein C, Bernstein H (September 1997). "Sexual communication". Journal of Theoretical Biology. 188 (1): 69–78. Bibcode:1997JThBi.188...69B. doi:10.1006/jtbi.1997.0459. PMID 9299310.
  25. ^ Danton H. O’Day, Paul A. Horgen (1981) Sexual Interactions in Eukaryotic Microbes Academic Press, New York. ISBN 0125241607 ISBN 978-0125241601
  26. ^ Dusenbery, David B. (2009). Living at Micro Scale, Chapters 19 & 20. Harvard University Press, Cambridge, Massachusetts ISBN 978-0-674-03116-6.
  27. ^ Raina AK, Klun JA (August 1984). "Brain factor control of sex pheromone production in the female corn earworm moth". Science. 225 (4661): 531–533. Bibcode:1984Sci...225..531R. doi:10.1126/science.225.4661.531. PMID 17750856. S2CID 40949867.
  28. ^ Xiang Y, Yang M, Li Z (2009). "Calling behavior and rhythms of sex pheromone production in the Black Cutworm Moth in China". Journal of Insect Behavior. 23 (1): 35–44. doi:10.1007/s10905-009-9193-0. S2CID 45577568.
  29. ^ Schulz S, Francke W, König WA, Schurig V, Mori K, Kittmann R, Schneider D (December 1990). "Male pheromone of swift moth, Hepialus hecta L. (Lepidoptera: Hepialidae)". Journal of Chemical Ecology. 16 (12): 3511–3521. Bibcode:1990JCEco..16.3511S. doi:10.1007/BF00982114. PMID 24263445. S2CID 26903035.
  30. ^ Papke RS, Kemp DJ, Rutowski RL (2007). "Multimodal Signalling: Structural Ultraviolet Reflectance Predicts Male Mating Success Better than Pheromones in the Butterfly Colias eurytheme L. (Pieridae)". Animal Behaviour. 73: 47–54. doi:10.1016/j.anbehav.2006.07.004. S2CID 40403665.
  31. ^ Burand JP, Tan W, Kim W, Nojima S, Roelofs W (2005). "Infection with the insect virus Hz-2v alters mating behavior and pheromone production in female Helicoverpa zea moths". Journal of Insect Science. 5: 6. doi:10.1093/jis/5.1.6. PMC 1283887. PMID 16299596.
  32. ^ Sen R, Gadagkar R (2010). "Natural history and behaviour of the primitively eusocial wasp (Hymenoptera: Vespidae): a comparison of the two sexes". Journal of Natural History. 44 (15–16): 959–968. doi:10.1080/00222931003615703. S2CID 84698285.
  33. ^ "Alpinobombus". Natural History Museum. Retrieved 26 September 2015
  34. ^ Svensson BG, Bergstrom G (1979). "Marking Pheromones of Alpinobornbus Males". Journal of Chemical Ecology. 5 (4): 603–615. Bibcode:1979JCEco...5..603S. doi:10.1007/bf00987845. S2CID 20759942.
  35. ^ Yao M, Rosenfeld J, Attridge S, Sidhu S, Aksenov V, Rollo CD (2009). "The Ancient Chemistry of Avoiding Risks of Predation and Disease". Evolutionary Biology. 36 (3): 267–281. Bibcode:2009EvBio..36..267Y. doi:10.1007/s11692-009-9069-4. ISSN 0071-3260. S2CID 29901266.
  36. ^ Schaal B, Coureaud G, Langlois D, Giniès C, Sémon E, Perrier G (July 2003). "Chemical and behavioural characterization of the rabbit mammary pheromone". Nature. 424 (6944): 68–72. Bibcode:2003Natur.424...68S. doi:10.1038/nature01739. PMID 12840760. S2CID 4428155.
  37. ^ Norris MJ (1954). "Sexual maturation in the desert locust, Schistocerca gregaria (Forskal), with special reference to the effects of grouping". Anti-Locust Bulletin. 18: 1–4.
  38. ^ a b c "Trace amine receptor: Introduction". International Union of Basic and Clinical Pharmacology. Archived from the original on 23 February 2014. Retrieved 15 February 2014. Importantly, three ligands identified activating mouse Taars are natural components of mouse urine, a major source of social cues in rodents. Mouse Taar4 recognizes β-phenylethylamine, a compound whose elevation in urine is correlated with increases in stress and stress responses in both rodents and humans. Both mouse Taar3 and Taar5 detect compounds (isoamylamine and trimethylamine, respectively) that are enriched in male versus female mouse urine. Isoamylamine in male urine is reported to act as a pheromone, accelerating puberty onset in female mice [34]. The authors suggest the Taar family has a chemosensory function that is distinct from odorant receptors with a role associated with the detection of social cues. ... The evolutionary pattern of the TAAR gene family is characterized by lineage-specific phylogenetic clustering [26,30,35]. These characteristics are very similar to those observed in the olfactory GPCRs and vomeronasal (V1R, V2R) GPCR gene families.
  39. ^ a b c d e f Liberles SD (October 2015). "Trace amine-associated receptors: ligands, neural circuits, and behaviors". Current Opinion in Neurobiology. 34: 1–7. doi:10.1016/j.conb.2015.01.001. PMC 4508243. PMID 25616211. Furthermore, while some TAARs detect aversive odors, TAAR-mediated behaviors can vary across species. ... The ability of particular TAARs to mediate aversion and attraction behavior provides an exciting opportunity for mechanistic unraveling of odor valence encoding.
    Figure 2: Table of ligands, expression patterns, and species-specific behavioral responses for each TAAR
  40. ^ a b c Wallrabenstein I, Singer M, Panten J, Hatt H, Gisselmann G (2015). "Timberol® Inhibits TAAR5-Mediated Responses to Trimethylamine and Influences the Olfactory Threshold in Humans". PLOS ONE. 10 (12): e0144704. Bibcode:2015PLoSO..1044704W. doi:10.1371/journal.pone.0144704. PMC 4684214. PMID 26684881. While mice produce gender-specific amounts of urinary TMA levels and were attracted by TMA, this odor is repellent to rats and aversive to humans [19], indicating that there must be species-specific functions. ... Furthermore, a homozygous knockout of murine TAAR5 abolished the attraction behavior to TMA [19]. Thus, it is concluded that TAAR5 itself is sufficient to mediate a behavioral response at least in mice. ... Whether the TAAR5 activation by TMA elicits specific behavioral output like avoidance behavior in humans still needs to be examined.
  41. ^ a b Pantages E, Dulac C (December 2000). "A novel family of candidate pheromone receptors in mammals". Neuron. 28 (3): 835–845. doi:10.1016/S0896-6273(00)00157-4. PMID 11163270.
  42. ^ Carlson NR (2013). Physiology of behavior (11th ed.). Boston: Pearson. p. 335. ISBN 978-0-205-23939-9.
  43. ^ Keverne EB (October 1999). "The vomeronasal organ". Science. 286 (5440): 716–720. doi:10.1126/science.286.5440.716. PMID 10531049.
  44. ^ Hildebrand JG (January 1995). "Analysis of chemical signals by nervous systems". Proceedings of the National Academy of Sciences of the United States of America. 92 (1): 67–74. Bibcode:1995PNAS...92...67H. doi:10.1073/pnas.92.1.67. PMC 42818. PMID 7816849.
  45. ^ Stoka AM (June 1999). "Phylogeny and evolution of chemical communication: an endocrine approach". Journal of Molecular Endocrinology. 22 (3): 207–225. doi:10.1677/jme.0.0220207. PMID 10343281.
  46. ^ R.S. Herz, T. Engen, Odor memory: review and analysis, Psychon. Bull. Rev. 3 (1996) 300–313.
  47. ^ Sherborne AL, Thom MD, Paterson S, Jury F, Ollier WE, Stockley P, et al. (December 2007). "The genetic basis of inbreeding avoidance in house mice". Current Biology. 17 (23): 2061–2066. Bibcode:2007CBio...17.2061S. doi:10.1016/j.cub.2007.10.041. PMC 2148465. PMID 17997307.
  48. ^ Jiménez JA, Hughes KA, Alaks G, Graham L, Lacy RC (October 1994). "An experimental study of inbreeding depression in a natural habitat". Science. 266 (5183): 271–273. Bibcode:1994Sci...266..271J. doi:10.1126/science.7939661. PMID 7939661.
  49. ^ a b c Foster RL (1992). "Nestmate Recognition as an Inbreeding Avoidance Mechanism in Bumble Bees (Hymenoptera: Apidae)". Journal of the Kansas Entomological Society. 65 (3): 238–243. JSTOR 25085362.
  50. ^ Martin SJ, Carruthers JM, Williams PH, Drijfhout FP (August 2010). "Host specific social parasites (Psithyrus) indicate chemical recognition system in bumblebees". Journal of Chemical Ecology. 36 (8): 855–863. Bibcode:2010JCEco..36..855M. doi:10.1007/s10886-010-9805-3. PMID 20509042. S2CID 4794525.
  51. ^ a b c d e Grammer K, Fink B, Neave N (February 2005). "Human pheromones and sexual attraction". European Journal of Obstetrics, Gynecology, and Reproductive Biology. 118 (2): 135–142. doi:10.1016/j.ejogrb.2004.08.010. PMID 15653193.
  52. ^ 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."
  53. ^ Hays WS (2003). "Human pheromones: have they been demonstrated?". Behavioral Ecology and Sociobiology. 54 (2): 89–97. doi:10.1007/s00265-003-0613-4. S2CID 37400635.
  54. ^ a b Bear MF, Connors BW, Paradiso MA (2006). Neuroscience: Exploring the Brain. Lippincott Williams & Wilkins. ISBN 978-0-7817-6003-4. neuroscience exploring the brain. p. 264 ...there has not yet been any hard evidence for human pheromones that might [change] sexual attraction (for members of either sex) [naturally]
  55. ^ Riley A (9 May 2016). "Pheromones are probably not why people find you attractive". BBC News. Retrieved 2016-05-09.
  56. ^ a b c d e f g Hays WS (2003). "Human pheromones: have they been demonstrated?". Behavioral Ecology and Sociobiology. 54 (2): 89–97. doi:10.1007/s00265-003-0613-4. S2CID 37400635.
  57. ^ a b c Mostafa T, El Khouly G, Hassan A (2012). "Pheromones in sex and reproduction: Do they have a role in humans?". Journal of Advanced Research. 3 (1): 1–9. doi:10.1016/j.jare.2011.03.003.
  58. ^ a b Kirk-Smith M (1978). "Human social attitudes affected by androstenol". Research Communications in Psychology, Psychiatry & Behavior. 3 (4): 379–384. ISSN 0362-2428.
  59. ^ McClintock MK (January 1971). "Menstrual synchorony and suppression". Nature. 229 (5282): 244–245. Bibcode:1971Natur.229..244M. doi:10.1038/229244a0. PMID 4994256. S2CID 4267390.
  60. ^ Stern K, McClintock MK (March 1998). "Regulation of ovulation by human pheromones". Nature. 392 (6672): 177–179. Bibcode:1998Natur.392..177S. doi:10.1038/32408. PMID 9515961. S2CID 4426700..
  61. ^ Yang Z, Schank JC (December 2006). "Women do not synchronize their menstrual cycles". Human Nature. 17 (4): 433–447. doi:10.1007/s12110-006-1005-z. PMID 26181612. S2CID 2316864.[permanent dead link]
  62. ^ Strassmann BI (March 1999). "Menstrual synchrony pheromones: cause for doubt". Human Reproduction. 14 (3): 579–580. doi:10.1093/humrep/14.3.579. PMID 10221677.
  63. ^ Van Toller C, Kirk-Smith M, Wood N, Lombard J, Dodd GH (1983). "Skin conductance and subjective assessments associated with the odour of 5-alpha-androstan-3-one". Biological Psychology. 16 (1–2): 85–107. doi:10.1016/0301-0511(83)90056-X. PMID 6682682. S2CID 54325922.
  64. ^ a b c Hummer TA, McClintock MK (April 2009). "Putative human pheromone androstadienone attunes the mind specifically to emotional information". Hormones and Behavior. 55 (4): 548–559. doi:10.1016/j.yhbeh.2009.01.002. PMID 19470369. S2CID 17022112.
  65. ^ Wyatt TD (April 2015). "The search for human pheromones: the lost decades and the necessity of returning to first principles". Proceedings. Biological Sciences. 282 (1804): 20142994. doi:10.1098/rspb.2014.2994. PMC 4375873. PMID 25740891.
  66. ^ a b c Michael RP, Bonsall RW, Kutner M (1975). "Volatile fatty acids, "copulins", in human vaginal secretions". Psychoneuroendocrinology. 1 (2): 153–163. doi:10.1016/0306-4530(75)90007-4. PMID 1234654. S2CID 38274482.
  67. ^ Liberles SD, Buck LB (August 2006). "A second class of chemosensory receptors in the olfactory epithelium". Nature. 442 (7103): 645–650. Bibcode:2006Natur.442..645L. doi:10.1038/nature05066. PMID 16878137. S2CID 2864195.
  68. ^ Pearson H (August 2006). "Mouse data hint at human pheromones". Nature. 442 (7102): 495. Bibcode:2006Natur.442..495P. doi:10.1038/442495a. PMID 16885951.
  69. ^ Wysocki CJ, Preti G (November 2004). "Facts, fallacies, fears, and frustrations with human pheromones". The Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology. 281 (1): 1201–1211. doi:10.1002/ar.a.20125. PMID 15470677.
  70. ^ Milinski M (2001). "Evidence for MHC-correlated perfume preferences in humans". Behavioral Ecology. 12 (2): 140–9. doi:10.1093/beheco/12.2.140.
  71. ^ Wedekind C, Seebeck T, Bettens F, Paepke AJ (June 1995). "MHC-dependent mate preferences in humans". Proceedings. Biological Sciences. 260 (1359): 245–249. Bibcode:1995RSPSB.260..245W. doi:10.1098/rspb.1995.0087. PMID 7630893. S2CID 34971350.
  72. ^ Purves D, Brannon EM, Cabeza R, LaBar KS, Huettel SA, Platt ML, Woldorff M (2008). Principles of Cognitive Neuroscience. Sinauer. ISBN 978-0-87893-694-6.

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