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Theory of creature signalling for evolutionary advantage

Within evolutionary biology, signalling theory is a body of theoretical work examining advice between individuals, both within species and across species. The central question is when organisms with alien interests, such as in sexual choice, should be expected to provide honest signals (no presumption being made of witting intention) rather than cheating. Mathematical models describe how signalling can contribute to an evolutionarily stable strategy.

Signals are given in contexts such as mate selection by females, which subjects the advertising males' signals to selective pressure. Signals thus evolve because they alter the behaviour of the receiver to benefit the signaller. Signals may be honest, conveying data which usefully increases the fitness of the receiver, or quack. An private tin can cheat by giving a dishonest signal, which might briefly benefit that signaller, at the run a risk of undermining the signalling system for the whole population.

The question of whether choice of signals works at the level of the individual organism or cistron, or at the level of the grouping, has been debated past biologists such as Richard Dawkins, arguing that individuals evolve to signal and to receive signals better, including resisting manipulation. Amotz Zahavi suggested that adulterous could be controlled by the handicap principle, where the all-time horse in a handicap race is the one carrying the largest handicap weight. According to Zahavi'due south theory, signallers such as male peacocks have 'tails' that are genuinely handicaps, beingness costly to produce. The organisation is evolutionarily stable as the large showy tails are honest signals. Biologists have attempted to verify the handicap principle, but with inconsistent results. The mathematical biologist Ronald Fisher analysed the contribution that having ii copies of each gene (diploidy) would brand to honest signalling, demonstrating that a runaway issue could occur in sexual selection. The evolutionary equilibrium depends sensitively on the balance of costs and benefits.

The same mechanisms tin be expected in humans, where researchers have studied behaviours including risk taking by young men, hunting of large game animals, and costly religious rituals, finding that these appear to qualify every bit plush honest signals.

Sexual selection [edit]

When animals choose mating partners, traits such as signalling are field of study to evolutionary pressure. For example, the male person gray tree frog, Hyla versicolor, produces a call to attract females. Once a female person chooses a mate, this selects for a specific style of male calling, thus propagating a specific signalling ability. The signal can exist the call itself, the intensity of a call, its variation way, its repetition rate, so on. Various hypotheses seek to explain why females would select for one telephone call over the other. The sensory exploitation hypothesis proposes that pre-existing preferences in female receivers can drive the evolution of signal innovation in male senders, in a similar way to the hidden preference hypothesis which proposes that successful calls are amend able to friction match some 'hidden preference' in the female person.[1] Signallers accept sometimes evolved multiple sexual ornaments,[2] and receivers have sometimes evolved multiple trait preferences.[3]

Honest signals [edit]

Eurasian jay, Garrulus glandarius, gives honest signals—loud alarm calls—from its tree perch when it sees a predator.

In biology, signals are traits, including structures and behaviours, that have evolved specifically because they change the behaviour of receivers in ways that benefit the signaller.[iv] Traits or actions that benefit the receiver exclusively are chosen "cues". For example, when an alarm bird deliberately gives a warning call to a stalking predator and the predator gives upward the chase, the audio is a "indicate". But when a foraging bird inadvertently makes a rustling audio in the leaves that attracts predators and increases the risk of predation, the sound is not a betoken, but a cue.[4]

Signalling systems are shaped by mutual interests between signallers and receivers. An alert bird such every bit a Eurasian jay warning off a stalking predator is communicating something useful to the predator: that it has been detected by the prey; it might as well quit wasting its time stalking this alerted prey, which it is unlikely to grab. When the predator gives upward, the signaller tin get back to other tasks such as feeding. Once the stalking predator is detected, the signalling prey and receiving predator thus have a mutual interest in terminating the chase.[5] [6]

Within species, mutual interests increase with kinship.[7] Kinship is central to models of signalling between relatives, for instance when broods of nestling birds beg and compete for nutrient from their parents.[eight] [9]

The term honesty in animal communication is controversial because in non-technical usage it implies intent, to discriminate deception from honesty in homo interactions.[6] Nevertheless, biologists use the phrase "honest signals" in a direct, statistical sense. Biological signals, like warning calls or resplendent tail feathers, are honest if they reliably convey useful information to the receiver. That is, the betoken trait[a] tells the receiver virtually an otherwise unobservable factor.[b] Honest biological signals practise not need to exist perfectly informative, reducing uncertainty to nil; all they need to be useful is to be right "on boilerplate", and then that some behavioural response to the signal is advantageous, statistically, compared to the behaviour that would occur in absence of the signal.[nine] Ultimately the value of the signalled information depends on the extent to which information technology allows the receiver to increase its fitness.[ten]

1 class of honest indicate is the aposematic warning point, generally visual, given by poisonous or dangerous animals such equally wasps, poison dart frogs, and pufferfish. Alert signals are honest indications of noxious prey, considering conspicuousness evolves in tandem with noxiousness (a conspicuous, non-noxious organism gets eaten). Thus, the brighter and more conspicuous the organism, the more toxic it usually is.[eleven] [12] The most common and effective colours are red, yellow, black and white.[13]

There is an obvious evolutionary reward to faking a signal similar "Don't swallow me, I'k poisonous"; the sender scares off predators without the metabolic cost of actually being toxic. Such false I'm-harmful signals are chosen Batesian mimicry. There is also an obvious evolutionary advantage to the predator learning to distinguish the real from the faux signal; this sets off an evolutionary artillery race, making such imitation signals evolutionarily unstable. An "honest signal", on the other hand, cannot exist faked by the sender (the feathers can't await resplendent unless the bird is good for you) or can't be faked without a toll that exceeds the benefit (like incessantly giving alert calls). This unfakability is what makes the signal reliable (and thus "honest"). Hence, "honest" signals are evolutionarily stable.

The mathematical biologist John Maynard Smith discusses whether honest signalling must ever be costly. He notes that it had been shown that "in some circumstances" a signal is reliable only if it is costly. He states that it had been assumed that parameters such every bit pay-offs and signalling costs were constant, but that this might be unrealistic. He states that with some restrictions, signals can be cost-gratuitous, reliable, and evolutionarily stable. However, if costs and benefits "vary uniformly over the whole range" then indeed honest signals have to be plush.[14]

Dishonest signals [edit]

Male fiddler crab signals with its enlarged fighting hook, just weak regrown claws may be dishonest signals.

Because in that location are both mutual and conflicting interests in near animal signalling systems, a key problem in signalling theory is dishonesty or cheating. For case, if foraging birds are safer when they give a warning call, cheats could requite simulated alarms at random, just in example a predator is nearby. But too much adulterous could crusade the signalling system to collapse. Every dishonest signal weakens the integrity of the signalling system, and so reduces the fettle of the group.[15] An example of quack signalling comes from Fiddler crabs such as Uca lactea mjoebergi, which accept been shown to bluff (no conscious intention beingness implied) nearly their fighting ability. When a claw is lost, a crab occasionally regrows a weaker claw that nevertheless intimidates crabs with smaller but stronger claws.[16] The proportion of dishonest signals is depression enough for it not to be worthwhile for venereal to test the honesty of every signal through combat.[xv]

Richard Dawkins and John Krebs in 1978 considered whether individuals of the same species would deed every bit if attempting to deceive each other. They applied a "selfish gene" view of development to animals' threat displays to see if it would be in their genes' interests to give dishonest signals. They criticised previous ethologists, such as Nikolaas Tinbergen and Desmond Morris for suggesting that such displays were "for the expert of the species". They argued that such communication ought to be viewed as an evolutionary arms race in which signallers evolve to go better at manipulating receivers, while receivers evolve to get more than resistant to manipulation.[fifteen] The game theoretical model of the war of attrition similarly suggests that threat displays ought non to convey whatever reliable data about intentions.[17]

Sports handicapping metaphor [edit]

The all-time horses in a handicap race carry the largest weights, and then the size of the handicap is a measure of the animal's quality.

In 1975, Amotz Zahavi proposed a verbal model for how signal costs could constrain cheating and stabilize an "honest" correlation between observed signals and unobservable qualities, based on an analogy to sports handicapping systems.[18] [19] He called this idea the handicap principle. The purpose of a sports handicapping arrangement is to reduce disparities in functioning, making the competition more competitive. In a handicap race, intrinsically faster horses are given heavier weights to comport under their saddles. Similarly, in amateur golf, better golfers have fewer strokes subtracted from their raw scores. This creates correlations betwixt the handicap and unhandicapped performance, if the handicaps work as they are supposed to, betwixt the handicap imposed and the corresponding horse'due south handicapped functioning. If nothing was known nigh two race horses or ii amateur golfers except their handicaps, an observer could infer who is most likely to win: the horse with the bigger weight handicap, and the golfer with the smaller stroke handicap. By analogy, if peacock 'tails' (large tail covert feathers) act as a handicapping system, and a peahen knew nothing about 2 peacocks except the sizes of their tails, she could "infer" that the peacock with the bigger tail has greater unobservable intrinsic quality. Display costs can include extrinsic social costs, in the form of testing and punishment by rivals, besides as intrinsic product costs.[20] Another example given in textbooks is the extinct Irish elk, Megaloceros giganteus. The male Irish elk's enormous antlers could mayhap have evolved as displays of ability to overcome handicap, though biologists point out that if the handicap is inherited, its genes ought to be selected against.[21]

Peacock signals reproductive fettle with its large colourful tail, possibly considering information technology is a handicap.

The essential idea here is intuitive and probably qualifies every bit folk wisdom. It was articulated by Kurt Vonnegut in his 1961 short story Harrison Bergeron.[22] In Vonnegut'due south futuristic dystopia, the Handicapper Full general uses a diverseness of handicapping mechanisms to reduce inequalities in performance. A spectator at a ballet comments: "information technology was easy to run across that she was the strongest and most graceful of all dancers, for her handicap bags were as big every bit those worn by 2 hundred pound men." Zahavi interpreted this analogy to hateful that higher quality peacocks with bigger tails are signalling their ability to "waste" more of some resources by trading it off for a bigger tail. This resonates with Thorstein Veblen's idea that conspicuous consumption and extravagant condition symbols tin betoken wealth.[23]

Zahavi'due south conclusions rest on his verbal estimation of a metaphor, and initially the handicap principle was not well received by evolutionary biologists.[19] However, in 1984, Nur and Hasson[24] used life history theory to show how differences in signalling costs, in the course of survival-reproduction tradeoffs, could stabilize a signalling arrangement roughly equally Zahavi imagined. Genetic models too suggested this was possible.[25] In 1990 Alan Grafen showed that a handicap-like signalling system was evolutionarily stable if college quality signallers paid lower marginal survival costs for their signals.[26]

In 1982, West.D. Hamilton proposed a specific only widely applicable handicap machinery, parasite-mediated sexual pick.[27] He argued that in the never-ending co-evolutionary race between hosts and their parasites, sexually selected signals indicate health. This thought was tested in 1994 in barn swallows, a species where males take long tail streamers. Møller found that the males with longer tails, and their offspring, did have fewer bloodsucking mites, whereas fostered young did not. The event was therefore genetic, confirming Hamilton's theory.[28]

Another example is Lozano's hypothesis that carotenoids have dual but mutually incompatible roles in immune function and signalling. Given that animals cannot synthesize carotenoids de novo, these must exist obtained from nutrient. The hypothesis states that animals with carotenoid-depended sexual signals are demonstrating their ability to "waste" carotenoids on sexual signals at the expense of their immune system.[29] [30]

The handicap principle has proven hard to examination empirically, partly because of inconsistent interpretations of Zahavi'southward metaphor and Grafen'southward marginal fitness model, and partly considering of conflicting empirical results: in some studies individuals with bigger signals seem to pay higher costs, in other studies they seem to be paying lower costs.[31] A possible explanation for the inconsistent empirical results is given in a series of papers by Getty,[32] [33] [6] [34] who shows that Grafen'south proof of the handicap principle is based on the critical simplifying assumption that signallers trade off costs for benefits in an additive style, the way humans invest money to increase income in the same currency.[c] Only the assumption that costs and benefits merchandise off in an additive fashion is truthful only on a logarithmic scale;[36] for the survival cost – reproduction do good tradeoff is assumed to mediate the evolution of sexually selected signals. Fitness depends on producing offspring, which is a multiplicative function of reproductive success given an private is still live times the probability of still being alive, given investment in signals.[24]

Costly signalling and Fisherian diploid dynamics [edit]

The try to detect how costs tin constrain an "honest" correlation between observable signals and unobservable qualities within signallers is built on strategic models of signalling games, with many simplifying assumptions. These models are virtually often applied to sexually selected signalling in diploid animals, but they rarely comprise a fact near diploid sexual reproduction noted by the mathematical biologist Ronald Fisher in the early 20th century: if there are "preference genes" correlated with choosiness in females as well as "signal genes" correlated with brandish traits in males, choosier females should tend to mate with showier males. Over generations, showier sons should too carry genes associated with choosier daughters, and choosier daughters should likewise carry genes associated with showier sons. This can cause the evolutionary dynamic known as Fisherian runaway, in which males become ever showier. Russell Lande explored this with a quantitative genetic model,[25] showing that Fisherian diploid dynamics are sensitive to signalling and search costs. Other models incorporate both costly signalling and Fisherian runaway.[37] [38] These models show that if fitness depends on both survival and reproduction, having sexy sons and choosy daughters (in the stereotypical model) can be adaptive, increasing fitness only as much as having healthy sons and daughters.[37] [38]

Examples [edit]

1 theory is that autumnal colours are a signal from copse to aphids of powerful chemical defences.

Sam Brownish and Due west. D. Hamilton, and Marco Archetti, proposed that autumn leaf colour is a betoken from copse to aphids and other pest species that migrate in autumn to the trees. In their theory, bright autumn coloration with pinks and yellows is costly to trees because pigments require free energy to synthesize, merely the investment may help them to reduce their parasite load.[39] [40]

Stotting, as in Thomson's gazelle, is cited as an example of signalling: the gazelles jump close to a predator instead of escaping, in what could exist a signal of strength.[41]

Man honest signals [edit]

Human behaviour may also provide examples of costly signals. In general, these signals provide data well-nigh a person'south phenotypic quality or cooperative tendencies. Evidence for plush signalling has been found in many areas of human interaction including chance taking, hunting, and religion.[42]

Costly signalling in hunting [edit]

A male hunter and a female gatherer of the Kali'na people of Guyana, fatigued by Pierre Barrère in 1743. Generous sharing by male hunters may serve as a "plush signal", helping them to acquire mates.

Large game hunting has been studied extensively equally a signal of men's willingness to accept concrete risks, as well as showcase force and coordination.[42] [43] [44] [45] Costly signalling theory is a useful tool for understanding food sharing amidst hunter gatherers considering it can be practical to situations in which delayed reciprocity is not a feasible caption.[46] [47] [48] Instances that are particularly inconsistent with the delayed reciprocity hypothesis are those in which a hunter shares his kill indiscriminately with all members of a large group.[49] In these situations, the individuals sharing meat accept no command over whether or non their generosity volition be reciprocated, and gratis riding becomes an attractive strategy for those receiving meat. Free riders are people who reap the benefits of grouping-living without contributing to its maintenance.[50] Fortunately, costly signalling theory can fill some of the gaps left past the delayed reciprocity hypothesis.[51] [52] Hawkes has suggested that men target large game and publicly share meat to draw social attention or to evidence off.[53] [48] Such brandish and the resulting favorable attention can improve a hunter'due south reputation by providing data most his phenotypic quality. Loftier quality signallers are more successful in acquiring mates and allies. Thus, costly signalling theory tin can explain apparently wasteful and altruistic behaviour.[18] [26] [52] [54] [55] [xviii] [56]

In order to be effective, plush signals must fulfill specific criteria.[18] [42] [57] Firstly, signallers must incur different levels of price and benefit for signalling behaviour. Secondly, costs and benefits must reflect the signallers' phenotypic quality. Thirdly, the information provided past a indicate should exist directed at and attainable to an audition. A receiver can be anyone who stands to do good from information the signaller is sending, such as potential mates, allies, or competitors. Honesty is guaranteed when but individuals of high quality tin pay the (loftier) costs of signalling. Hence, costly signals make it incommunicable for low-quality individuals to fake a signal and fool a receiver.[eighteen] [42] [57]

Bliege Bird et al. observed turtle hunting and spear line-fishing patterns in a Meriam customs in the Torres Strait of Australia, publishing their findings in 2001. Here, just some Meriam men were able to accrue high caloric gains for the amount of time spent turtle hunting or spear fishing (reaching a threshold measured in kcal/h). Since a daily catch of fish is carried dwelling house by hand and turtles are oftentimes served at large feasts, members of the community know which men near reliably brought them turtle meat and fish. Thus, turtle hunting qualifies as a costly betoken. Furthermore, turtle hunting and spear line-fishing are actually less productive (in kcal/h) than foraging for shellfish, where success depends merely on the amount of time dedicated to searching, and so shellfish foraging is a poor signal of skill or strength. This suggests that energetic gains are non the primary reason men have part in turtle hunting and spear angling.[42] A follow-up study found that successful Meriam hunters practice feel greater social benefits and reproductive success than less skilled hunters.[58]

The Hadza people of Tanzania also share food, possibly to gain in reputation.[59] Hunters cannot be sharing meat mainly to provision their families or to gain reciprocal benefits, every bit teenage boys often give away their meat even though they do non nonetheless have wives or children, so costly signalling of their qualities is the likely explanation.[lx] These qualities include skillful eyesight, coordination, forcefulness, cognition, endurance, or bravery. Hadza hunters more often pair with highly fertile, hard-working wives than non-hunters.[57] A woman benefits from mating with a man who possesses such qualities as her children will most likely inherit qualities that increment fitness and survivorship. She may also benefit from her husband'southward loftier social condition. Thus, hunting is an honest and costly signal of phenotypic quality.[52]

Among the men of Ifaluk atoll, costly signalling theory tin also explain why men torch fish.[61] Torch fishing is a ritualized method of fishing on Ifaluk whereby men use torches made from dried coconut fronds to catch large canis familiaris-toothed tuna. Preparation for torch fishing requires significant time investments and involves a smashing deal of organization. Due to the fourth dimension and energetic costs of preparation, torch fishing results in internet caloric losses for fishers. Therefore, torch fishing is a handicap that serves to signal men's productivity.[61] Torch line-fishing is the nearly advertised fishing occupation on Ifaluk. Women and others usually spend time observing the canoes as they sheet beyond the reef. Too, local rituals assistance to broadcast data about which fishers are successful and enhance fishers' reputations during the torch fishing season. Several ritual behavioural and dietary constraints clearly distinguish torch fishers from other men. First, males are simply permitted to torch fish if they participated on the first twenty-four hours of the line-fishing season. The community is well informed equally to who participates on this day, and can easily identify the torch fishers. 2d, torch fishers receive all of their meals at the canoe house and are prohibited from eating certain foods. People frequently discuss the qualities of torch fishermen. On Ifaluk, women claim that they are looking for hard-working mates.[62] With the singled-out sexual segmentation of labor on Ifaluk, industriousness is a highly valued characteristic in males.[63] Torch fishing thus provides women with reliable data on the piece of work ethic of prospective mates, which makes information technology an honest costly point.[52]

In many human cases, a strong reputation built through plush signalling enhances a human being's social status over the statuses of men who betoken less successfully.[49] [64] [65] Amid northern Kalahari foraging groups, traditional hunters unremarkably capture a maximum of 2 or iii antelopes per year.[66] Information technology was said of a peculiarly successful hunter:[67]

"It was said of him that he never returned from a hunt without having killed at to the lowest degree a wildebeest, if not something larger. Hence the people connected with him ate a keen deal of meat and his popularity grew."[67]

Although this hunter was sharing meat, he was not doing so in the framework of reciprocity.[67] The general model of costly signalling is not reciprocal; rather, individuals who share learn more mates and allies.[18] [42] Costly signalling applies to situations in Kalahari foraging groups where giving often goes to recipients who have little to offering in render. A immature hunter is motivated to print customs members with daughters so that he can obtain his first married woman. Older hunters may wish to concenter women interested in an extramarital human relationship, or to be a co-wife.[68] [69] In these northern Kalahari groups, the killing of a large animate being indicates a man who has mastered the art of hunting and tin can support a family. Many women seek a homo who is a good hunter, has an agreeable graphic symbol, is generous, and has advantageous social ties.[66] [69] [seventy] Since hunting ability is a prerequisite for marriage, men who are good hunters enter the wedlock marketplace earliest. Costly signalling theory explains seemingly wasteful foraging displays.[57]

Concrete risks every bit a costly indicate [edit]

Young men may have role in risky sports like motorcycle racing to bespeak their forcefulness and skill.

Costly signalling can be practical to situations involving concrete strain and risk of physical injury or decease.[42] [71] Research on physical adventure taking is important because information regarding why people, especially young men, accept part in loftier chance activities tin help in the development of prevention programs.[72] [71] Reckless driving is a lethal trouble among young men in western societies.[72] A male who takes a physical run a risk is sending the bulletin that he has plenty strength and skill to survive extremely unsafe activities. This indicate is directed at peers and potential mates.[eighteen] When those peers are criminals or gang members, sociologists Diego Gambetta and James Densley detect that hazard-taking signals can assist expedite credence into the grouping.[73] [74]

In a study of take chances taking, some types of hazard, such every bit concrete or heroic risk for others' benefit, are viewed more favorably than other types of risk, such as taking drugs. Males and females valued different degrees of heroic take chances for mates and aforementioned-sexual activity friends. Males valued heroic gamble taking by male friends, just preferred less of it in female person mates. Females valued heroic risk taking in male mates and less of it in female friends. Females may exist attracted to males inclined to physically defend them and their children. Males may prefer heroic risk taking by male person friends as they could be adept allies.[71]

In western societies, voluntary claret donation is a common, all the same less extreme, course of run a risk taking. Costs associated with these donations include pain and hazard of infection.[75] If blood donation is an opportunity to send costly signals, then donors volition be perceived by others as generous and physically salubrious.[18] [76] In a survey, both donors and non-donors expressed perceptions of the wellness, generosity, and ability of blood donors to operate in stressful situations.[76]

Religion as a costly betoken [edit]

Religious rituals such as snake handling may be explainable as costly signals.

Costly religious rituals such as male and female person circumcision, food and h2o deprivation, and snake handling expect paradoxical in evolutionary terms. Devout religious beliefs wherein such traditions are proficient appear maladaptive.[77] Religion may take arisen to increase and maintain intragroup cooperation.[78] Cooperation leads to altruistic behaviour,[79] and costly signalling could explain this.[xviii] All religions may involve costly and elaborate rituals, performed publicly, to demonstrate loyalty to the religious group.[80] In this way, grouping members increment their allegiance to the group by signalling their investment in group interests. Nonetheless, as group size increases amidst humans, the threat of free riders grows.[l] Plush signalling theory accounts for this by proposing that these religious rituals are costly enough to deter free riders.[81]

Irons proposed that costly signalling theory could explain plush religious behaviour. He argued that difficult-to-simulated religious displays enhanced trust and solidarity in a customs, producing emotional and economic benefits. He showed that display signals among the Yomut Turkmen of northern Iran helped to secure trade agreements. These "ostentatious" displays signalled commitment to Islam to strangers and group members.[82] Sosis demonstrated that people in religious communities are four times more probable to live longer than their secular counterparts,[51] [79] and that these longer lifespans were positively correlated with the number of costly requirements demanded from religious community members.[83] However, confounding variables may not take been excluded.[84] Wood found that religion offers a subjective feeling of well-being within a community, where costly signalling protects confronting gratis riders and helps to build self-command among committed members.[85] Iannaccone studied the effects of costly signals on religious communities. In a cocky-reported survey, every bit the strictness of a church increased, the attendance and contributions to that church increased proportionally. In result, people were more than willing to participate in a church that has more stringent demands on its members.[81] Despite this observation, costly donations and acts conducted in a religious context does not itself establish that membership in these clubs is actually worth the entry costs imposed.

Despite the experimental back up for this hypothesis, it remains controversial. A common critique is that devoutness is like shooting fish in a barrel to false, such as simply by attending a religious service.[86] Even so, the hypothesis predicts that people are more likely to join and contribute to a religious grouping when its rituals are plush.[81] Another critique specifically asks: why religion? There is no evolutionary reward to evolving religion over other signals of commitment such every bit nationality, as Irons admits. However, the reinforcement of religious rites likewise every bit the intrinsic reward and penalty system establish in organized religion makes it an ideal candidate for increasing intragroup cooperation. Finally, there is insufficient show for increase in fitness as a consequence of religious cooperation.[79] However, Sosis argues for benefits from religion itself, such as increased longevity, improved health, aid during crises, and greater psychological well-being,[87] although both the supposed benefits from religion and the plush-signaling mechanism have been contested.[88]

Run across also [edit]

  • Alarm bespeak
  • Conspicuous consumption
  • Dramaturgy (sociology)
  • Game theory
  • Greenish-beard upshot
  • Origin of language
  • Signalling (economics)
  • Virtue signalling
  • Zoosemiotics

Notes [edit]

  1. ^ Economists telephone call what is bachelor to the receiver "public information".
  2. ^ Economists call the unobservable affair that would exist of value to the receiver "private information"; biologists often phone call it "quality"
  3. ^ Grafen'due south proof is formally similar to a classic monograph on economical marketplace signalling by Nobel laureate Michael Spence.[35]

References [edit]

  1. ^ Gerhardt, Humfeld & Marshall 2007.
  2. ^ Møller & Pomiankowski 1993.
  3. ^ Pomiankowski & Iwasa 1993.
  4. ^ a b Bradbury & Vehrenkamp 1998.
  5. ^ Bergstrom & Lachmann 2001.
  6. ^ a b c Getty 2002.
  7. ^ Johnstone 1998.
  8. ^ Godfray 1995.
  9. ^ a b Johnstone 1999.
  10. ^ Dall et al. 2005.
  11. ^ Maan & Cummings 2012.
  12. ^ Blount et al. 2009.
  13. ^ Stevens & Ruxton 2012.
  14. ^ Maynard Smith 1994.
  15. ^ a b c Dawkins & Krebs 1978.
  16. ^ Lailvaux, Reaney & Backwell 2009.
  17. ^ Caryl 1979.
  18. ^ a b c d east f g h i Zahavi 1975.
  19. ^ a b Zahavi 1997.
  20. ^ Searcy & Nowicki 2005.
  21. ^ Feldhamer 2007, p. 423.
  22. ^ Vonnegut 1961.
  23. ^ Veblen 1899.
  24. ^ a b Nur & Hasson 1984.
  25. ^ a b McElreath & Boyd 2007.
  26. ^ a b Grafen 1990.
  27. ^ Hamilton & Zuk 1982.
  28. ^ Møller 1994.
  29. ^ Lozano 1994.
  30. ^ McGraw & Ardia 2003.
  31. ^ Kotiaho 2001.
  32. ^ Getty 1998a.
  33. ^ Getty 1998b.
  34. ^ Getty 2006.
  35. ^ Spence 1974.
  36. ^ Tazzyman, Iwasa & Pomiankowski 2014.
  37. ^ a b Eshel, Sansone & Jacobs 2002.
  38. ^ a b Kokko 2002.
  39. ^ Hamilton & Dark-brown 2001.
  40. ^ Archetti 2000.
  41. ^ Maynard Smith & Harper 2003.
  42. ^ a b c d e f g Bliege Bird, Smith & Bird 2001.
  43. ^ Gurven & Hill 2009.
  44. ^ Hawkes 1990.
  45. ^ Wiessner 2002.
  46. ^ Bliege Bird & Bird 1997.
  47. ^ Gurven et al. 2000.
  48. ^ a b Hawkes 1993.
  49. ^ a b Wiessner 1996.
  50. ^ a b Barrett, Dunbar & Lycett 2002.
  51. ^ a b Sosis 2000b.
  52. ^ a b c d Smith & Bliege Bird 2000.
  53. ^ Hawkes 1991.
  54. ^ Johnstone 1995.
  55. ^ Johnstone 1997.
  56. ^ Zahavi 1977.
  57. ^ a b c d Hawkes & Bliege Bird 2002.
  58. ^ Smith, Bliege Bird & Bird 2002.
  59. ^ Marlowe 2010.
  60. ^ Hawkes, O'Connell & Blurton_Jones 2001.
  61. ^ a b Sosis 2000a.
  62. ^ Sosis, Feldstein & Hill 1998.
  63. ^ Sosis 1997.
  64. ^ Kelly 1995.
  65. ^ Dowling 1968.
  66. ^ a b Lee 1979.
  67. ^ a b c Thomas 1959.
  68. ^ Lee 1993.
  69. ^ a b Shostak 1981.
  70. ^ Marshall 1976.
  71. ^ a b c Farthing 2005.
  72. ^ a b Nell 2002.
  73. ^ Densley 2012.
  74. ^ Gambetta 2009.
  75. ^ Schreiber et al. 2006.
  76. ^ a b Lyle, Smith & Sullivan 2009.
  77. ^ Tuzin 1982.
  78. ^ Steadman & Palmer 2008.
  79. ^ a b c Bulbulia 2004.
  80. ^ Irons 2001.
  81. ^ a b c Iannaccone 1992.
  82. ^ Irons 1996.
  83. ^ Sosis & Bressler 2003.
  84. ^ Hood, Hill & Spilka 2009.
  85. ^ Forest 2016.
  86. ^ Rees 2009.
  87. ^ Sosis 2003.
  88. ^ Schuurmans-Stekhoven 2016.

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Further reading [edit]

  • Zahavi, Amotz (1977). "The cost of honesty (Farther remarks on the handicap principle)". Journal of Theoretical Biology. 67 (three): 603–605. Bibcode:1977JThBi..67..603Z. doi:10.1016/0022-5193(77)90061-iii. PMID 904334.
  • Zahavi, Amotz (1977). "The Testing of the Bond". Animal Behaviour. 25: 246–247. doi:10.1016/0003-3472(77)90089-6. S2CID 53197593.

External links [edit]

  • Brute behavior online: Cant

waddingtonuntoonesch.blogspot.com

Source: https://en.wikipedia.org/wiki/Signalling_theory

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