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SHETTLEWORTH - AnBehav 2000 - Animal Cognition and Behavior
SHETTLEWORTH - AnBehav 2000 - Animal Cognition and Behavior
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Animal cognition and animal behaviour
SARA J. SHETTLEWORTH
Cognitive processes such as perception, learning, memory and decision making play an important role in
mate choice, foraging and many other behaviours. In this review, I summarize a few key ideas about
animal cognition developed in a recent book (Shettleworth 1998, Cognition, Evolution and Behaviour) and
briefly review some areas in which interdisciplinary research on animal cognition is currently proving
especially productive. Cognition, broadly defined, includes all ways in which animals take in information
through the senses, process, retain and decide to act on it. Studying animal cognition does not entail any
particular position on whether or to what degree animals are conscious. Neither does it entail rejecting
behaviourism in that one of the greatest challenges in studing animal cognition is to formulate clear
behavioural criteria for inferring specific mental processes. Tests of whether or not apparently goaldirected behaviour is controlled by a representation of its goal, episodic-like memory in birds, and
deceptive behaviour in monkeys provide examples. Functional modelling has been integrated with
analyses of cognitive mechanisms in a number of areas, including studies of communication, models of
how predator learning and attention affect the evolution of conspicuous and cryptic prey, tests of the
relationship betweeen ecological demands on spatial cognition and brain evolution, and in research on
social learning. Rather than a new field of cognitive ecology, such interdisciplinary research on animal
cognition exemplifies a revival of interest in proximate mechanisms of behaviour.
Correspondence: S. J. Shettleworth, Department of Psychology, University of Toronto, 100 St George Street, Toronto, Ontario M5S 3G3,
Canada (email: shettle@psych.utoronto.ca).
00033472/01/020277+10 $35.00/0
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depth of an interdisciplinary approach to communication is abundantly evident in two recent books (Hauser
1996; Bradbury & Vehrencamp 1998). In other areas
too, prominent research programmes have successfully
integrated psychological and biological thinking about
cognitive mechanisms. Some of them are outlined below.
In this article, I first discuss a few general issues in the
study of animal cognition. Some of these are areas where
misunderstandings or gaps in communication still seem
to exist. Then I briefly review some areas where interdisciplinary research on cognition is proving especially
illuminating. In a recent book (Shettleworth 1998), I
argued at length that the most rewarding way toward
understanding animal cognition is to integrate data and
theory from biology and psychology. I tried to contribute
to such an integration by juxtaposing different kinds of
studies of animal information use that are not always
considered together and to point out what they have
in common and what unanswered questions and new
theoretical perspectives such a juxtapostion generates. I
particularly hoped to help students of animal cognitive
processes coming from different backgrounds to appreciate the richness of what fields other than their own have
to offer and to provide them with an entre e into what
may be alien literatures. Over the years, others have made
similar arguments (e.g. Rozin & Schull 1988; Stamps
1991; Yoerg 1991; Kamil 1988, 1998). Indeed, over 30
years ago Hindes (1966) influential textbook of animal
behaviour was subtitled A Synthesis of Ethology and
Comparative Psychology.
WHAT IS COGNITION?
Cognition, broadly defined, includes perception, learning,
memory and decision making, in short all ways in which
animals take in information about the world through the
senses, process, retain and decide to act on it. Like most
working definitions, this notion is soft around the edges,
but it has the merit of encouraging a comprehensive study
of animal information processing. Authors (e.g. Tomasello
& Call 1997) sometimes write as if animal behaviour can
be divided into the noncognitive or reflexive and the
cognitive, which usually means behaviour more complex
or flexible than anything thought to be explicable by
simple reflexes or associations. However, as discussed
below, even simple associative learning can result in complex representational structures that allow a degree of
flexibility sometimes attributed to higher learning. It is
usually impossible to know in advance of detailed experimental analysis what kind of process underlies behaviour.
For instance, alarm calling and responding to alarms may
be a reflex or, as outlined below, it may be mediated by a
representation of the type of predator being signalled and
modulated by contextual information like the proximity
of kin (Holmes & Sherman 1983). A comprehensive study
of comparative cognition that embraces all of these
mechanisms invites one to analyse the function and evolutionary history of different levels of complexity within a
single theoretical framework.
Except in the context of cognitive ethology (Griffin
1978, 1998; Ristau 1991), the study of animal cognition is
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Cognitive Ecology?
The biology and psychology of cognition are being
synthesized in a number of other areas. Flower constancy
in bees has been elegantly related to detailed properties of
memory (Chittka et al. 1999). The adaptive value of
learning has been investigated by showing that growth or
reproduction are greater in experimental environments
where important resources like mates or oviposition sites
can be associated with simple cues than in environments
where the same resources are unpredictable (see Dukas &
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CONCLUSIONS
The possible benefits of integrating mechanistic and
functional, or psychological and biological, approaches
to animal cognition have been discussed many times (e.g.
Shettleworth 1984, 1998; Kamil 1988; Rozin & Schull
1988; Stamps 1991). Nevertheless, some of the barriers to
interdisciplinary communication pointed out by Kamil &
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Yoerg (1982) still seem to exist. Among them are differences in terminology and in attitudes to laboratory versus
field research among workers from different backgrounds.
Thinking in terms of proximal causes does not always
come easily to people accustomed to thinking in terms of
ultimate causes and vice versa, but such intellectual
flexibility can surely be facilitated by more widespread
interdisciplinary training. An earlier review of learning
and behavioural ecology (Shettleworth 1984) could
point to only a few cases, mostly from foraging, in which
any aspect of cognition was being studied in its ecological
context. Developments in behavioural ecology, ethology
and comparative cognition in the ensuing years have
contributed to the much richer menu of cognitive mechanisms and functional problems sampled in this article.
There is every indication that this richness is still
increasing.
Acknowledgments
Preparation of this article was supported by a Research
Grant from the Natural Sciences and Engineering
Research Council of Canada. I thank Rob Hampton, Marc
Hauser, Sue Healy, Ed Wasserman, David Sherry and two
anonymous referees for comments on the manscript.
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