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Within
Signals are given in contexts such as mate selection by females, which subjects the males' signals to selective pressure. Signals thus evolve because they modify the behaviour of the receiver to benefit the signaller. Signals may be honest, conveying information which usefully increases the fitness of the receiver, or dishonest. An individual can cheat by giving a dishonest signal, which might briefly benefit that signaller, at the risk of undermining the signalling system for the whole population.
The question of whether selection of signals works at the level of the individual organism or gene, or at the level of the group, has been debated by biologists such as
The same mechanisms can 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 as costly honest signals.
Sexual selection
When animals choose mates,
Honest signals
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.[2] Traits or actions that benefit the receiver exclusively are called cues. When an alert bird deliberately gives a warning call to a stalking predator and the predator gives up the hunt, the sound is a signal. When a foraging bird inadvertently makes a rustling sound in the leaves that attracts predators and increases the risk of predation, the sound is a 'cue'.[2]
Signalling systems are shaped by mutual interests between signallers and receivers. An alert bird such as 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 catch. When the predator gives up, the signaller can 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 hunt.[3][4]
Within species, mutual interests increase with kinship.[5] Kinship is central to models of signalling between relatives, for instance when broods of nestling birds beg and compete for food from their parents.[6][7]
The concept of honesty in animal communication is controversial because it is difficult to determine intent and use that as a criterion to discriminate deception from honesty, as we do in human interactions.[8] Biologists use the phrase "honest signals" in a statistical sense. Biological signals, like warning calls or resplendent tail feathers, are honest if they are correlated with something useful to the receiver, that is, the signal trait[a] is correlated with the unobservable thing of value to the receiver.[b] Honest biological signals do not need to be perfectly informative, reducing uncertainty to zero; all they need to be useful is to be correct "on average".[9] Ultimately the value of the signalled information depends on the extent to which it allows the receiver to increase its fitness.[10] Hence, "honest" signals are evolutionarily stable.
![](http://upload.wikimedia.org/wikipedia/commons/thumb/b/b8/Yellow-banded.poison.dart.frog.arp.jpg/170px-Yellow-banded.poison.dart.frog.arp.jpg)
One class of honest signal is the warning signal (aposematism), generally visual, given by poisonous or dangerous animals such as the yellow-banded and other poison dart frogs. Warning signals are honest indications of noxious prey, because conspicuousness evolves in tandem with noxiousness.[11] Thus, the brighter and more conspicuous the organism, the more toxic it usually is.[11] The most common and effective colours are red, yellow, black and white.[12]
Dishonest signals
![](http://upload.wikimedia.org/wikipedia/commons/thumb/3/31/Uca_lactea-male-600-450.jpg/220px-Uca_lactea-male-600-450.jpg)
Because there are both mutual and conflicting interests in most animal signalling systems, a central problem in signalling theory is dishonesty or
The sports handicapping metaphor
![](http://upload.wikimedia.org/wikipedia/commons/thumb/b/b7/Curlin_in_the_Final_Turn_in_the_2008_Stephen_Foster_Handicap_%282972085393%29.jpg/300px-Curlin_in_the_Final_Turn_in_the_2008_Stephen_Foster_Handicap_%282972085393%29.jpg)
In 1975,
The essential idea here is intuitive and probably qualifies as
Zahavi’s conclusions rest on his verbal interpretation 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 form of survival-reproduction tradeoffs, could stabilize a signalling system roughly as Zahavi imagined. Genetic models also suggested this was possible.[25] In 1990 Alan Grafen showed that a handicap-like signalling system was evolutionarily stable if higher quality signallers paid lower marginal survival costs for their signals.[26]
In 1982,
Another example is Lozano's hypothesis that
The handicap principle has proven hard to test empirically, partly because of inconsistent interpretations of Zahavi’s metaphor and Grafen’s marginal fitness model, and partly because 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][32] A possible explanation for the inconsistent empirical results is given in a series of papers by Getty,[33][34][35][36] who shows that Grafen’s proof of the handicap principle is based on the critical simplifying assumption that signallers trade off costs for benefits in an additive fashion, the way humans invest money to increase income in the same currency.[c] But the assumption that costs and benefits trade off in an additive fashion is not valid for the survival cost – reproduction benefit tradeoff that 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 individual is still alive times the probability of still being alive, given investment in signals.[24]
Costly signalling and Fisherian diploid dynamics
The effort to discover how costs can 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 most often applied to
Examples
![](http://upload.wikimedia.org/wikipedia/commons/thumb/9/92/Autumn.westonbirt.750pix.jpg/220px-Autumn.westonbirt.750pix.jpg)
Sam Brown and
Human honest signals
Human behaviour may also provide examples of costly signals. In general, these signals provide information about a person’s phenotypic quality or cooperative tendencies. Evidence for costly signalling has been found in many areas of human interaction including risk taking, hunting, and religion,[43] which are discussed in turn below.
Costly signalling in hunting
![](http://upload.wikimedia.org/wikipedia/commons/thumb/c/cc/Kalina_hunter_gatherer.jpg/220px-Kalina_hunter_gatherer.jpg)
Large game hunting has been studied extensively as a signal of men’s willingness to take physical risks, as well as showcase strength and coordination.
In order to be effective, costly signals must fulfill specific criteria.[18][43][58] Firstly, signallers must incur different levels of cost and benefit for signalling behaviour. Secondly, costs and benefits must reflect the signallers’ phenotypic quality. Thirdly, the information provided by a signal should be directed at and accessible to an audience. A receiver can be anyone who stands to benefit from information the signaller is sending, such as potential mates, allies, or competitors. Honesty is guaranteed when only individuals of high quality can pay the (high) costs of signalling. Hence, honest signalling make it impossible for low-quality individuals to fake a signal and fool a receiver.[18][43][58]
Bliege Bird et al. observed turtle hunting and spear fishing patterns in a Meriam community in the Torres Strait of Australia, publishing their findings in 2001. Here, only some Meriam men were able to accumulate 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 home by hand and turtles are frequently served at large feasts, members of the community know which men most reliably brought them turtle meat and fish. Thus, turtle hunting qualifies as a costly signal. Furthermore, turtle hunting and spear fishing are actually less productive (in kcal/h) than foraging for shellfish, where success depends only on the amount of time dedicated to searching, so shellfish foraging is a poor signal of skill or strength. This suggests that energetic gains are not the primary reason men take part in turtle hunting and spear fishing.[43] A follow-up study found that successful Meriam hunters do experience greater social benefits and reproductive success than less skilled hunters.[59]
The Hadza people of Tanzania also share food, possibly to gain in reputation.[60] Hunters cannot be sharing meat mainly to provision their families or to gain reciprocal benefits, as teenage boys often give away their meat even though they do not yet have wives or children, so costly signalling of their qualities is the likely explanation.[61] These qualities include good eyesight, coordination, strength, knowledge, endurance, or bravery. Hadza hunters more often pair with highly fertile, hard-working wives than non-hunters.[62] A woman benefits from mating with a man who possesses such qualities as her children will most likely inherit qualities that increase fitness and survivorship. She may also benefit from her husband’s high social status. Thus, hunting is an honest and costly signal of phenotypic quality.[63]
Among the men of
In many human cases, a strong reputation built through costly signalling enhances a man’s social status over the statuses of men who signal less successfully.
- "It was said of him that he never returned from a hunt without having killed at least a wildebeest, if not something larger. Hence the people connected with him ate a great deal of meat and his popularity grew."[69]
Although this hunter was sharing meat, he was not doing so in the framework of reciprocity.[69] The general model of costly signalling is not reciprocal; rather, individuals who share acquire more mates and allies.[18][43] Costly signalling applies to situations in Kalahari foraging groups where giving often goes to recipients who have little to offer in return. A young hunter is motivated to impress community members with daughters so that he can obtain his first wife. Older hunters may wish to attract women interested in an extramarital relationship, or to be a co-wife.[70][71] In these northern Kalahari groups, the killing of a large animal indicates a man who has mastered the art of hunting and can support a family.[72] Generally, many women seek a man who is a good hunter, has an agreeable character, is generous, and has advantageous social ties.[68][71][72] Since hunting ability is a prerequisite for marriage, men who are good hunters enter the marriage market earliest. Costly signalling theory explains seemingly wasteful foraging displays.[62]
Physical risks as a costly signal
![](http://upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Motorcycle_racing.jpg/225px-Motorcycle_racing.jpg)
Costly signalling can be applied to situations involving physical strain and risk of physical injury or death.[73][74][75] Research on physical risk taking is important because information regarding why people, especially young men, take part in high risk activities can help in the development of prevention programs.[74][75] Reckless driving is a lethal problem among young men in western societies.[76] A male who takes a physical risk is sending the message that he has enough strength and skill to survive extremely dangerous activities. This signal is directed at peers and potential mates.[18]
In a study of risk taking, some types of risk, such as physical 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 risk for mates and same-sex friends. Males valued heroic risk taking by male friends, but preferred less of it in female mates. Females valued heroic risk taking in male mates and less of it in female friends. Females may be attracted to males inclined to physically defend them and their children. Males may prefer heroic risk taking by male friends as they could be good allies.[75]
In western societies, voluntary blood donation is a common, yet less extreme, form of risk taking. Costs associated with these donations include pain and risk of infection.[77] If blood donation is an opportunity to send costly signals, then donors will be perceived by others as generous and physically healthy.[18][78] In a survey, both donors and non-donors expressed perceptions of the health, generosity, and ability of blood donors to operate in stressful situations.[78]
Religion as a costly signal
![](http://upload.wikimedia.org/wikipedia/commons/thumb/4/4b/Handling_of_serpents%2C_a_part_of_the_ceremony_at_the_Pentecostal_Church_of_God._This_coal_camp_offers_none_of_the..._-_NARA_-_541340.jpg/170px-Handling_of_serpents%2C_a_part_of_the_ceremony_at_the_Pentecostal_Church_of_God._This_coal_camp_offers_none_of_the..._-_NARA_-_541340.jpg)
Costly religious rituals such as male circumcision, food and water deprivation, and snake handling look paradoxical in evolutionary terms. Devout religious beliefs wherein such traditions are practiced therefore appear maladaptive.[79] Religion may have arisen to increase and maintain intragroup cooperation.[80] Cooperation leads to altruistic behaviour,[81] and costly signalling could explain this.[18] All religions may involve costly and elaborate rituals, performed publicly, to demonstrate loyalty to the religious group.[82] In this way, group members increase their allegiance to the group by signalling their investment in group interests. However, as group size increases among humans, the threat of free riders grows.[51] Costly signalling theory accounts for this by proposing that these religious rituals are costly enough to deter free riders.[83]
Irons proposed that costly signalling theory could explain costly religious behaviour. He argued that hard-to-fake religious displays enhanced trust and solidarity in a community, producing emotional and economic benefits. He showed that display signals among the
Despite the experimental support for this hypothesis, it remains controversial. A common critique is that devoutness is easy to fake, such as simply by attending a religious service.[88] However, the hypothesis predicts that people are more likely to join and contribute to a religious group when its rituals are costly.[87] Another critique specifically asks: why religion? There is no evolutionary advantage to evolving religion over other signals of commitment such as nationality, as Irons admits.[81] However, the reinforcement of religious rites as well as the intrinsic reward and punishment system found in religion makes it an ideal candidate for increasing intragroup cooperation. Finally, there is insufficient evidence for increase in fitness as a result of religious cooperation.[81] However, Sosis argues for benefits from religion itself, such as increased longevity, improved health, assistance during crises, and greater psychological well being.[89]
See also
- Animal communication
- Autumn leaf color
- Baculum
- Cheap talk
- Conspicuous consumption
- Dramaturgy (sociology)
- Handicap principle
- Multiple sexual ornaments
- Origin of language
- Signalling (economics)
- Signalling game
Notes
- ^ Economists call what is available to the receiver "public information".
- ^ Economists call the unobservable thing that would be of value to the receiver "private information"; biologists often call it "quality"
- ^ Grafen’s proof is formally similar to a classic monograph on economic market signalling by Nobel laureate Michael Spence.[37]
References
- ^ Gerhardt et al, 2007
- ^ a b Bradbury 1998
- ^ Bergstrom 2001
- ^ Getty 2002
- ^ Johnstone 1998
- ^ Godfray 1995
- ^ Johnstone 1999
- ^ Getty 1997
- ^ Johnstone 1993
- ^ Dall 2005
- ^ JSTOR 663197.
- .
- ^ a b c Krebs and Dawkins 1982
- ^ a b Dawkins and Krebs 1978
- ^ BES 2008
- ^ Lailvaux 2008
- ^ Caryl, 1979
- ^ a b c d e f g Zahavi 1975
- ^ a b Zahavi 1997
- ^ Searcy 2005
- ^ Feldhamer, 2007. p. 423.
- ^ Vonnegut 1961
- ^ Veblen 1899
- ^ a b Nur 1984
- ^ a b McElreath 2007
- ^ a b Grafen 1990
- ^ Hamilton 1982
- ^ Møller 1994
- ^ a b Lozano 1994
- ^ McGraw 2003
- ^ Møller 1999
- ^ Kotiaho 2001
- ^ Getty 1998a
- ^ Getty 1998b
- ^ Getty 2002a
- ^ Getty 2006
- ^ Spence 1974
- ^ a b Eshel 2002
- ^ a b Kokko 2002
- ^ a b Hamilton 2001
- ^ a b Archetti 2000
- ^ Maynard Smith 2003
- ^ a b c d e f Bliege Bird et al. 2001
- ^ Gurven and Hill 2009
- ^ Hawkes 1990
- ^ Weissner 2002
- ^ Bliege Bird and Bird 1997
- ^ Gurven et al. 2000
- ^ Hawkes 1993
- ^ a b Wiessner 1996
- ^ a b Barrett et al. 2002
- ^ a b c d Sosis 2000
- ^ a b c Smith and Bliege Bird 2000
- ^ Hawkes (1991, 1993)
- ^ Getty 1998
- ^ Johnstone 1995, 1997
- ^ Zahavi 1975, 1977
- ^ a b Hawkes and Bliege Bird 2002
- ^ Smith et al. 2002
- ^ Marlowe 2010
- ^ Hawkes 2001
- ^ a b Hawkes et al. 2002
- ^ Smith et al. 2000
- ^ Sosis et al. 1998
- ^ Sosis 1997
- ^ Kelly 1995
- ^ Dowling 1968
- ^ a b Lee 1979
- ^ a b c Thomas 1959
- ^ Lee 1993
- ^ a b Shostak 1981
- ^ a b Marshall 1976
- ^ Bleige Bird et al. 2001
- ^ a b Nell 2002
- ^ a b c Farthing 2005
- ^ Nell 2001
- ^ Schreiber et al. 2006
- ^ a b Lyle et al. 2009
- ^ Tuzin 1982
- ^ Steadman and Palmer 2008
- ^ a b c d Bulbulia 2004
- ^ Irons 2001
- ^ Iannaccone 1992
- ^ Irons 1996
- ^ Sosis and Bressler 2003
- ^ Hood et al. 2009
- ^ a b Iannacone 1992
- ^ Rees 2011
- ^ Sosis 2002
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{{cite book}}
: CS1 maint: multiple names: authors list (link) - Zahavi A (1977) Reliability in communication systems and the evolution of altruism. In: Stonehouse B, Perrins CM (eds) Evolutionary ecology. Macmillan, London, 253– 259.
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External links
- An Introduction to the Theory of Honest Signalling
- Animal behavior online: Deceit
- Journal of Management: Signaling Theory: A Review and Assessment (2011, behind paywall)
Category:Animal communication Category:Evolutionary biology