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MARC BEKOFF AND PAUL W. SHERMAN 4 Reflections on Animal Selves IS SELF-COGNIZANCE a uniquely human attribute, or do other animals also have a sense of self? Although there is considerable interest in this question, answers remain elusive. Progress has been stymied by misunderstandings in terminology, a focus on a narrow range of species, and controversies over key concepts, experimental paradigms and interpretations of data. Here, we propose a new conceptual and terminological framework, emphasizing that degrees of self-cognizance differ among animals because of the cognitive demands that their species-specific social structures and life-history characteristics have placed upon them over evolutionary time. We suggest that the self-cognizance of an organism falls at a point on a continuum of social complexity and conscious involvement. Although laypersons and researchers from many disciplines have long been interested in animal self-knowledge (or self-cognizance) [1–3], and a large amount of information has accumulated, few unambiguous conclusions are available. This is mainly because of the difficulty of objectively assessing selfknowledge and quantifying its neurobiological substrates among organisms whose patterns of communication we only partially understand. Progress has also been limited by inconsistencies in terminology, and by experimental paradigms that concentrate on visual rather than on chemical or auditory cues. Moreover, relatively few species have been examined in detail, and all were vertebrates , mostly primates. Here, we suggest that it is appropriate and useful to consider knowledge of self, or ‘self-cognizance’, as a continuum ranging from self-referencing to self-awareness to self-consciousness (Box 1). We argue that degrees of selfcognizance are better predicted by the behavioral ecology of a species rather than by its relative brain size or phylogenetic closeness to humans. In social animals, cognitive demands imposed by selection for cooperation, maintenance of pair bonds, nepotism, and reciprocity on the one hand, and avoidance of being cheated and effectiveness in competition on the other hand, have resulted in the evolution of increased mental complexity [1–3]. Studies of selfcognizance will benefit from capitalizing on this diversity, and also from considering self-cognizance in invertebrates and vertebrates. We argue that the degree of self-cognizance of individuals in any species can be represented as a point on a continuum of complexity and conscious involvement. Reprinted from Bekoff, M., and P. W. Sherman. 2004. Reflections on Animal Selves. Trends in Ecology and Evolution 19, 176–180, with permission from Elsevier. However, documenting degrees of self-cognizance is difficult. Ideally, individuals should be studied in their natural environments when they are making decisions about how to modify their behavior toward other individuals of their social group in light of the previous responses of those group members to them. Thus, it will be useful to combine field observations of dynamic changes in the behavior of an individual in social situations requiring selfcognizance , such as deciding how long or hard to fight over a resource [4] or responding to being cheated in a social contract [5,6], with noninvasive neural techniques to determine whether the target behaviors are linked to electrophysiological responses of the types, and in the specific brain regions, that are active in self-cognizant humans. Currently, technical difficulties preclude applying certain neural techniques to field situations. However, relevant techniques are being developed and, in the meantime, cleverly designed laboratory experiments [5,7,8] might enable Reflections on Animal Selves 67 BOX 1. CATEGORIES (DEGREES) OF SELF-COGNIZANCE In our scheme, ‘self-cognizance’ is used as an umbrella term to cover the continuum from self-referencing to self-consciousness. We hypothesize that species exhibit different degrees of self-cognizance, which reflect variations in their social environments and life histories. The position of an individual on the selfcognizance continuum is based on the degree to which members of its species or group engage in repetitive competitive or cooperative interactions with the same conspecifics over their lifetimes and benefit from changing their responses in light of outcomes of those previous interactions (see also [1,7,23]). Self-Referencing (also referred to as self-referent phenotype matching and the ‘armpit effect’: [17,31,42–45]). Self-referencing is a perceptual process involving matching phenotypic characteristics of a target individual against the phenotype of the discriminator . Discriminators compare labels of the target (such as of odor or appearance) against labels learned from their own phenotype, and accept or reject that target based on the degree of similarity [42,46]. Self-referencing can be reflexive and noncognitive, even occurring in the immune system...


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