PSY 2364 Animal Communication. Tinbergen s four questions. Niko Tinbergen ( ) Niko Tinbergen ( ) Niko Tinbergen ( )

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PSY 2364 Animal Communication The curious naturalist noninvasive field experiments Studies of herring gull behavior, spatial learning in insects Tinbergen s four questions 1) What is the mechanism? 2) What is its function? 3) How did it develop (ontogeny)? 4) How did it evolve (phylogeny)? Tinbergen, N. 1963. On aims and methods of ethology. Zeitschrift für Tierpsychologie 20: 410-433 Search image: a hypothetical mental picture of a prey item used by a predator to search specifically for a cryptic, common, edible prey. Red-tailed hawk Sign stimulus the component of an action or object that triggers a fixed response in an animal (e.g., herring gull chick s begging response) Sign stimulus the component of an action or object that triggers a fixed response in an animal (e.g., herring gull chick s begging response) Red dot 1

Innate or learned? Innate or learned? Jack Hailman (1967) studied the begging response of herring gull chicks to two different models of adult birds: herring gull models and laughing gull models. Herring gull model Laughing gull model Hailman found that newly hatched herring gulls do not distinguish between the two models. However, at 6 days, they can reliably tell the difference. Releaser a sign stimulus given by one animal to another as a social signal (e.g. pheromones chemical communication signals) Yawning A Releaser? White-tailed deer alarm signal Konrad Lorenz (1903-1989) Studied how innate and acquired components of behavior are integrated Emphasized the importance of behavior for taxonomy Studied imprinting in geese Origins and basis of animal and human aggression Konrad Lorenz (1903-1989) Fixed action pattern innate, stereotyped response triggered by a well-defined simple stimulus. Once activated, the response is always performed to completion. 2

Konrad Lorenz (1903-1989) Imprinting form of learning in which individuals exposed to certain key stimuli, usually during an early stage of development, form an association with the object and may later show sexual behavior toward similar objects. Konrad Lorenz (1903-1989) Konrad Lorenz Konrad Lorenz (1903-1989) Critical period for learning a period in the life span of an individual (in birds, a short period just after hatching) where learning or imprinting is greatly facilitated. Fixed action pattern: an innate, highly stereotyped response triggered by a welldefined simple stimulus; once activated, the response is performed in its entirety. Egg rolling in geese Lorenz & Tinbergen (1939) https://www.youtube.com/watch?v=vunzv-bypku Graylag Goose http://www.youtube.com/watch?v=vunzv-bypku 3

Instinct: a behavior pattern that reliably develops in individuals that receive adequate nutrition, and that is given in functional form on its first performance. Honesty and communication Information conveyed by animal signals can provide information about the environment or the sender s intentions (future course of action). Honesty and communication Most (but not all) communication signals are honest (reliable). Why is this? Zahavi (1975) argued that receivers will not respond (and hence communication will not evolve) if the signal is dishonest or unreliable. Honesty and communication Handicap hypothesis: some signals, like the peacock s tail, are costly to produce. Since they are costly, they are more likely to be reliable (and this is why female peacocks prefer them). Zahavi argued that costly signals are more likely to evolve; he describes these as handicaps. Intention Movements Ethologists proposed that some communication signals originate from movements made when the animal is getting ready to perform a particular behavior. These movements are called intention movements. Examples: bared teeth display in dogs; upright threat posture in herring gulls 4

Displacement Behavior Ethologists observed that animals may sometimes engage in irrelevant activities when they experience conflict. They called these redirected activities displacement behaviors. Examples: cat grooms itself when prevented from reaching food; courtship begging in birds Ritualization Evolutionary process that transforms an incidental cue/behavior pattern into a true communication signal (Tinbergen, 1952). Simplification or reduction in components Exaggeration of the remaining components Repetition of the signal Stereotypy reduction in the signal variance e.g. courtship feeding in pheasants Evolution of communication signals Charles Darwin (1809-1882) Natural selection results in genetic change in a population when individuals differ in heritable traits that are correlated with differences in their reproductive success Principle of antithesis Darwin proposed that some communication signals arise (evolve) because animals are divided between conflicting tendencies or impulses (e.g. fear and aggression). Darwin s principle of antithesis states that signals with opposite messages tend to be opposite in form. Antithesis Opposition between two conflicting emotional states (attack vs. flee) Displays often show extremes in posture, orientation, extent of feather erection in birds, or fur in mammals start Male three-spined sticklebacks in breeding condition will perform a zig-zag dance in the presence of a female, but a different display (head-down threat posture) when approached by another male. 5

Tinbergen (1951) showed that the cue evoking the zig-zag courtship display was the swollen belly of the female fish; threat was induced by the red color on the underside of a rival male. Tinbergen found that simple (inanimate) fish models were sufficient to trigger the response (even a red post office van driving past the window near the fish tank!) Sign stimuli adapted for communication are called releasers. Specialization for signal production Specialization for signal perception discrimination vs. generalization innate releasing mechanisms lock-andkey analogy Template and filter models Egg recognition in herring gulls (Baerends) Supernormal stimuli Code breaking The relationship between the triggering cue and the behavioral response of a FAP can be observed when the behavior is exploited by members of another, parasitic species. Code breaking: behavior that mimics the triggering component of a FAP to exploit its benefits. Code breaking Examples of code breaking colored gape of cuckoo nestling parasitic rove beetle larva mimic the foodbegging behavior of larval ants Mock DW. (1999) Driving parents cuckoo. Nature 397, 647-648 6

Supernormal stimuli preference for giant eggs by Oystercatchers Supernormal stimuli supernormal begging behavior in brood parasites (European cuckoo, brown-headed cowbirds)? 7