Signalling Theory

peacock

Within evolutionary biology, signalling theory is a body of theoretical work examining communication between individuals. The central question is when organisms with conflicting interests should be expected to communicate honestly (no presumption being made of conscious intention) rather than cheating. Mathematical models in which organisms signal their condition to other individuals as part of an evolutionarily stable strategy are important for research in this field.

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, on average conveying information that is actually useful to the receiver, increasing its fitness, or dishonest. A cheat can give a dishonest signal, gaining an advantage, but potentially undermining the signalling system for all.

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 Richard Dawkins, arguing that individuals evolve to signal and to receive signals better, including resisting manipulation. Amotz Zahavi suggested that cheating could be controlled by the handicap principle, which suggests that reliable signals must be costly to the signaler, costing the signaler something that could not be afforded by an individual with less of a particular trait. For example, in the case of sexual selection, the theory suggests that animals of greater biological fitness signal this status through handicapping behavior or morphology that effectively lowers this quality.

The central idea is that sexually selected traits function like conspicuous consumption, signalling the ability to afford to squander a resource. Receivers know that the signal indicates quality because inferior quality signalers cannot afford to produce such wastefully extravagant signals. 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.

When animals choose mates, traits such as signalling are subject to evolutionary pressure. For example, the male gray tree frog produces a call to attract females. Once a female chooses a mate, this selects for a specific style of male calling, thus propagating a specific signalling ability. The signal can be the call itself, the intensity of a call, its variation style, its repetition rate, and so on. Various hypotheses seek to explain why females would select for one 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 better able to match some ‘hidden preference’ in the female.

In biology, signals are traits, including structures and behaviours, that have evolved specifically because they change the behavior of receivers in ways that benefit the signaller. 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.’

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 terminating the hunt. Within species, mutual interests increase with kinship. Kinship is central to models of signalling between relatives, for instance when broods of nestling birds beg and compete for food from their parents.

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. 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. Honest biological signals do not need to be perfectly informative, reducing uncertainty to zero; they only need to be correct ‘on average’ to be useful. Ultimately the value of the signalled information depends on the extent to which it allows the receiver to increase its fitness. Hence, ‘honest’ signals are evolutionarily stable.

Because there are both mutual and conflicting interests in most animal signalling systems, a central problem in signalling theory is dishonesty or cheating. For example, if foraging birds are safer when they give a warning call, cheats could give false alarms at random, just in case a predator is nearby. But too much cheating could cause the signalling system to collapse. Every dishonest signal weakens the integrity of the signalling system, and so reduces the fitness of the group. An example of dishonest signalling comes from Fiddler crabs, which have been shown to bluff (no conscious intention being implied) about their fighting ability. When a claw is lost, a crab occasionally regrows a weaker claw that nevertheless intimidates crabs with smaller but stronger claws. The proportion of dishonest signals is low enough for it not to be worthwhile for crabs to test the honesty of every signal through combat.

Richard Dawkins and John Krebs in 1978 considered whether individuals of the same species would act as if attempting to deceive each other. They applied a ‘selfish gene’ view of evolution to animals’ threat displays to see if it would be in their genes’ interests to give dishonest signals. They criticized previous ethologists, such as Nikolaas Tinbergen and Desmond Morris for suggesting that such displays were ‘for the good of the species.’ They argued that such communication ought to be viewed as an evolutionary arms race in which signallers evolve to become better at manipulating receivers, while receivers evolve to become more resistant to manipulation. The game theoretical model of the war of attrition similarly suggests that threat displays ought not to convey any reliable information about intentions.

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 sexually selected signalling in diploid animals (organisms with paired chromosomes), but they rarely incorporate a fact about 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 display traits in males, choosier females should tend to mate with showier males. Over generations, showier sons should also carry genes associated with choosier daughters, and choosier daughters should also carry genes associated with showier sons. This can cause the evolutionary dynamic known as ‘Fisherian runaway,’ in which males become ever showier. This model shows that if fitness depends on both survival and reproduction, having sexy sons and choosy daughters can be adaptive, increasing fitness just as much as having healthy sons and daughters.

Human behavior 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. 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. Costly signalling theory is a useful tool for understanding food sharing among hunter gatherers because it can be applied to situations in which delayed reciprocity is not a viable explanation. 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.

In these situations, the individuals sharing meat have no control over whether or not their generosity will be reciprocated, and free riding becomes an attractive strategy for those receiving meat. Free riders are people who reap the benefits of group-living without contributing to its maintenance. Fortunately, Costly signalling theory can fill some of the gaps left by the delayed reciprocity hypothesis. Hawkes has suggested that men target large game and publicly share meat to draw social attention or to show off. Such display and the resulting favorable attention can improve a hunter’s reputation by providing information about his phenotypic quality. High quality signallers are more successful in acquiring mates and allies. Thus, costly signalling theory can explain apparently wasteful and altruistic behavior.

Bliege Bird et al. observed turtle hunting and spearfishing 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. 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 spearfishing are actually less productive 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 spearfishing. A follow-up study found that successful Meriam hunters do experience greater social benefits and reproductive success than less skilled hunters.

Among the men of Ifaluk atoll, costly signalling theory can also explain why men torch fish. Torch fishing is a ritualized method of fishing on Ifaluk whereby men use torches made from dried coconut fronds to catch large dog-toothed tuna. Preparation for torch fishing requires significant time investments and involves a great deal of organization. Due to the time and energetic costs of preparation, torch fishing results in net caloric losses for fishers. Therefore, torch fishing is a handicap that serves to signal men’s productivity. Torch fishing is the most advertised fishing occupation on Ifaluk. Women and others usually spend time observing the canoes as they sail beyond the reef. Also, local rituals help to broadcast information 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 only permitted to torch fish if they participated on the first day of the fishing season. The community is well informed as to who participates on this day, and can easily identify the torch fishers. Second, 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 hardworking mates. With the distinct sexual division of labor on Ifaluk, industriousness is a highly valued characteristic in males. Torch fishing thus provides women with reliable information on the work ethic of prospective mates, which makes it an honest costly signal.

Costly signalling can be applied to situations involving physical strain and risk of physical injury or death. 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. Reckless driving is a lethal problem among young men in western societies. 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.

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.

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. Religion may have arisen to increase and maintain intragroup cooperation. Cooperation leads to altruistic behavior, and costly signalling could explain this. All religions may involve costly and elaborate rituals, performed publicly, to demonstrate loyalty to the religious group. 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. Costly signalling theory accounts for this by proposing that these religious rituals are costly enough to deter free riders.

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