Terkel conducted an in-depth study aimed to determine whether the observed behavior, the systematic stripping of pine cone scales from pine cones prior to eating, was a socially acquired behavior, as this action had not been observed elsewhere. The experimentation with and observation of these black rats was one of the first to integrate field observations with laboratory experiments to analyze the social learning involved.
Animal migration may be in part cultural; released ungulates have to learn over generations the seasonal changes in local vegetation.
In the black rat Rattus rattus , social transmission appears to be the mechanism of how optimal foraging techniques are transmitted. Terkel et al. Only young offspring could learn the technique. This result suggests that this optimal foraging technique is socially rather than genetically transmitted. Birds have been a strong study subject on the topic of culture due to their observed vocal "dialects" similar to those studied in the cetaceans. These dialects were first discovered by zoologist Peter Marler , who noted the geographic variation in the songs of various songbirds.
However, despite this hindrance, evidence for differing dialects among songbird populations has been discovered, especially in sparrows , starlings , and cowbirds. In these birds, scientists have found strong evidence for imitation-based learning, one of the main types of social learning. Though the songbirds obviously learn their songs through imitating other birds, many scientists remain skeptical about the correlation between this and culture: " It is how imitation affects and is affected by context, by ongoing social behavior, that must be studied before assuming its explanatory power.
Located in the eastern and southern parts of North America, Zonotrichia leucophrys is a species of white-crowned song-birds that exhibit learned vocal behavior. In this experimental setting, male birds in acoustic chambers were exposed to recorded sounds played through a loudspeaker.
It was also shown that white-crowned sparrows selectively learn songs of conspecifics. However, social bond strongly facilitates song imitation in some songbirds. Zebra finches hardly imitate songs played from a loudspeaker, but they can imitate songs of an adult bird after only a few hours of interaction . Interestingly, imitation is inhibited when the number of siblings pupils increases .
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Thus, the imitation process cannot be explained as simple mimicry that depends on the abundance of the model, but is a dynamic process that depends on complex behavioral context. In the British Isles, in the 20th century, bottled milk was delivered to households in the early morning by "milkmen" and left on doorsteps to be collected. Birds, particularly but not exclusively tits Paridae and principally blue tit Cyanistes caeruleus, great tit Parus major, coal tit Periparus ater regularly attacked the bottles, opening the foil or cardboard lids, and drank the cream of the top.
This behaviour was studied, and strong evidence found that learning was involved, although possibly at the level that milk bottles represent a food source, rather than particular techniques. Later studies built on these observations see the references in the two papers cited. Evidence for cultural transmission has also been shown in wild fish populations. Scientists Helfman and Schultz  conducted translocation experiments with French grunts Haemulon flavolineatum where they took fish native to a specific schooling site and transported them to other sites.
In this species of fish, the organism uses distinct, traditional migration routes to travel to schooling sites on coral reefs. These routes persisted past one generation and so by relocating the fish to different sites, Helfman and Schultz wanted to see if the new fish could relearn that sites' migration route from the resident fish. Indeed this is what they found: that the newcomers quickly learned the traditional routes and schooling sites. But when residents were removed under similar situations, the new fish did not use the traditional route and instead use new routes, suggesting that the behavior could not be transmitted once the opportunity for learning was no longer there.
In a similar experiment looking at mating sites in blueheaded wrasse Thalassoma bifasciatum , researcher Warner found that individuals chose mating sites based on social traditions and not based on the resource quality of the site. Warner found that although mating sites were maintained for four generations, when entire local populations were translocated elsewhere, new sites were used and maintained.
A popular method of approaching the study of animal culture and its transmission is the "ethnographic method," which argues that culture causes the geographical differences in the behavioral repertoires of large-brained mammals. However, this significantly downplays the roles that ecology and genetics play in influencing behavioral variation from population to population within a species.
Behaviors stemming from genetic or environmental effects are not reliant on socially learned and transmitted information; therefore, they are not cultural. Culture is just one source of adaptive behavior an organism exhibits to better exploit its environment. When behavioral variation reflects differential phenotypic plasticity , it is due more to ecological pressures than cultural ones. In other words, when an animal changes its behavior over its lifespan, this is most often a result of changes in its environment.
Furthermore, animal behavior is also influenced by evolved predispositions, or genetics. It is very possible that " correlation between distance between sites and 'cultural difference' might reflect the well-established correlation between genetic and geographical distances".
Another argument against the "ethnographic method" is that it is impossible to prove that there are absolutely no ecological or genetic factors in any behavior. However, this criticism can also be applied to studies of human culture. Though culture has long been thought to arise and remain independent of genetics, the constraints on the propagation and innovation of cultural techniques inevitably caused by the genome of each respective animal species has led to the theory of gene-culture coevolution , which asserts that "cognitive, affective, and moral capacities" are the product of an evolutionary dynamic involving interactions between genes and culture over extended periods of time.
We do not know every possible genetic or environmental effect on behavior that exists, nor will we ever. In other words, it is impossible to reject the notion that genes and ecology influence all behaviors, to a degree. Culture can also contribute to differences in behavior, but like genes and environments, it carries different weight in different behaviors.
As Laland and Janik  explain, "to identify cultural variation, not only is it not sufficient to rule out the possibility that the variation in behavior constitutes unlearned responses to different selection pressures [from the environment], but it is also necessary to consider the possibility of genetic variation precipitating different patterns of learning.
Some scientists believe the study of animal culture should be approached in a different way.
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Therefore it seems as though scientists should focus on examining how much variance in a behavior can be attributed to culture. Performing field experiments is an excellent way to try to answer this question: translocating individuals between populations or populations between sites could help biologists distinguish between culture, ecology and genetics. For example, if a newly introduced animal shifts its behavior to mimic that of others in its new population, genetic difference as an effect on behavior can be ruled out. In the study of social transmissions, one of the important unanswered questions is an explanation of how and why maladaptive social traditions are maintained.
In this case, not only is the social tradition maladaptive, but it also inhibits the acquisition of adaptive behavior. From Wikipedia, the free encyclopedia. This article is about learning in non-human animals. For the place of animals in human culture, see Animals in culture. Main article: Social learning in animals.
Animals portal. Animal cognition Animal communication Animal language Animal consciousness Animal-made art Animal training Culture Culture theory Cultural transmission Social learning in animals. New York: Basic Books, Galef, eds. The Question of Animal Culture. Cambridge, Massachusetts: Harvard UP, Laland and Vincent M. Organization Science. Culture and Cognition.
"It would almost be more surprising if these were simply random interactions."
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Trends in Cognitive Sciences. Cognitive Development in Chimpanzees. Tokyo: Springer, Adaptive Behavior 7: Animal Behaviour. The Quarterly Review of Biology. Lessons from Animal Teaching. Culture in Whales and Dolphins. The Behavioral and Brain Sciences. Davis; J. Krebs; S. West An Introduction to Behavioural Ecology 4th ed. Developments in primatology. Current Directions in Psychological Science. Whiten; A. Spiteri; V. Horner; K. Bonnie; S. Lambeth; S. Schapiro; F. Current Biology.