[Social learning in animals: comparative analysis of different forms and levels]

Social learning as one of the key concepts of cognitive ecology includes different forms of behavioural displays from relatively simple, such as "social release" and "stimulus enhancement" up to "teaching" and "cultural transmission" in animal communities. Rapid development of this fields resulted in some contradictions in methods and terminology. In this review different forms and levels of social learning are analyzed. Ecological aspects of social learning are connected with diet shaping, fear of predators and mate choice. The first aspect is the most studied but still discussible. Social learning being an intricate component of feeding behaviour matches with innate behaviour, imprinting as well as early associative learning. Investigation of cognitive aspects of social learning going back to Thorndike's crucial question "Do apes ape?" are now developing into series of questions including even: "Do ants ape?". Elaboration of universal methods of comparative studying of social learning such as "artificial fruit" and "two ways/one outcome" has essentially enlightened these questions and made comparative analysis possible. Large continuum of displays of cognitive skills in social learning has been revealed in non-primate species. One of the discussible issues in the role of social learning is distribution of innovations. Many authors have investigated this intriguing aspect of animal behaviour in different ways, such as long field observations as well as laboratory experiments based on "artificial innovators" that is specimens specially taught by experimentalists. Many impressive results were obtained; in particular it turned out in contradiction with some mathematical models that individuals in groups are rather different in their psychophysiological predisposition to innovative behaviour. Role of teaching in such different forms of behaviour as shaping of species-specific behavioural patterns and spread of innovations is considered. Although the majority of animals in wild populations are not good teachers and pupils, some cultural aspects of behaviour were recently revealed, mostly in primates. At the same time some classical results concerning cultural transmission of new patterns (for example, bottle-opening in tits) were experimentally revised. Many problems still remain unsolved, in particular, how spread of innovations may favour prosperity of populations; to what degree behavioural peculiarities of local groups may be determined by processes of social learning; which internal and external factors and under what circumstances invest into social learning in natural environment.