Neuron Review Primate Social Cognition: Uniquely Primate, Uniquely Social, or Just Unique? Richard W. Byrne 1, * and Lucy A. Bates 1 1 Centre for Social Learning and Cognitive Evolution and Scottish Primate Research Group, University of St Andrews, St Andrews KY16 9JP, Scotland, UK *Correspondence: [email protected]DOI 10.1016/j.neuron.2010.03 .010 Primates undoubtedly have impressive abilities in perceiving, recognizing, manipulating, and predicting other individuals, but only great apes seem to recognize the cognitive basis of manipulative and cooperative tactics or the concept of self. None of these abilities is unique to primates. We distinguish (1) a package ofquantitative advantages in social sophistication, perhaps based on more efficient memory, in which neocor- tical enlargement is associated with the challenge of social living; from (2) a qualitative difference in under- standing, whose taxonomic distr ibuti on—in cludin g sever al dista ntly relat ed specie s, inclu ding bird s— does not point to an evolu ti onar y or igin in soci al chall enges and may inst ead relate to a need to acquir e novel ways of dealing with the physical world. The ability of great apes to learn new manual routines by parsing action components may have driven their qualitatively greater social skill, suggesting that strict partition of physical and social cognition is likely to be misleading. Introduction Fift y years ago, human expe rimen tal psych olog y was dominated by learning theory: but the ‘‘cognitive revolution’’ dramatically chan ged the natur e of rese arch into human behavio r. Since then, neuroscientists have regularly used nonhuman primates as a ‘‘simpler’’ model to test theories of human social cognition and informati on proce ssing , with most laborat ory stud ies ofprimates relying on a few species of macaque monkeys. During the same period, a growing number of psychologists, anthropol- ogists, and ethologists began studying the social behavior ofa wid e ran ge of nonhuman primat es,both in theprima tes ’ natural habitat and in the laboratory, in a parallel attempt to reconstruct the evolutionary history of human mental competence. In this review, we shall attempt to explain what these wider- ranging studies have shown us about nonhuman primate social cognition, in the hope that it will inform understanding in neuro- science and new work on humans. It is important to note that there are numerous differences in behavior between different primate species, even among different species of monkeys— as well as between monkeys and great apes, of which humans are one of several extant species, including orangutans, gorillas, and chimp anze es. Thesedifferen ces may reflec t varia tions in the cognitive architecture, or they may result from socioecological differences between species that limit the expression of similar underlying cognitive capacity. Where appropriate, we therefore identify the primate species studied, in order to avoid the trap of treating all monkey species, or even all nonhuman primates, as alike in their behavior and cognition. Mi mi cking the paradi gm shif t in human expe ri ment al psychology, there has been a later and more gradual change towa rd cog nitiv e expl anati ons of animal beha vior.Earlierdebates about behavior were often dominated by the issue of whether ‘‘ simpl e’ ’ asso ciat ive learn ing coul d acco unt for the data or whether the animals ‘‘were cognitive,’’ usually taken to mean having conscious thought processes (Byrne and Bates, 2006; Macp hail, 1998 ) . As in the ca se of human e xp e ri m en ta l psychology, the associative approach was not refuted in prin- ciple, but the sheer effort and increasingly ad hoc appearance of associationist explanations for complex behavior led to the ascendance of cogn itive theoriz ing. A parti cular impetus for change came from the growing evidence of sophistication and comple xit y in pri mate beh avi or: in tak ing acc oun t of a ric h netwo rk of relat ionsh ips (Chene y and Seyfarth , 1982a , 1986, 1990 ), in inter preti ng othe rs’ intentions (Menz el, 1971, 1974; Premack, 1988; Premack and Woodruff, 1978 ), and in deploying subtl e manipulati ve tactics (Byr ne and Whiten, 1985, 1991, 1992; de Waal, 1982, 1986; Whiten and Byrne, 1988 ). By the mid-19 90s, it had become acc ept abl e to compar e across species by taki ng a gene rallycogniti ve appr oach, frame d in terms of infor matio n processing rathe r than cons ciousnes s (Byrne, 1995; Tomase llo and Call , 1997 ). This shift toward cognit ive explanations of behavior has now left us much better placed to gene rate and test hypoth eses about the evol utio n of human cognitive skill. Much of the new evidence of complexity in primate behavior came from the soci al real m, and comp arab le sophi stic atio n was not known for physical cognition (Byrne and Whiten, 1988; Cheney and Seyfarth, 1985; Jolly, 1966 ). This pattern meshed with Humphrey’s influential theory (Humphrey, 1976, 1981 ) that the evol utio nary chal leng e for which the solut ion is ‘‘ intel lige nce’ ’ is more likely a social than a physical one. He argued that semi- permanent social living, as found in many primate species, sets up a selective pressure for increasing social sophistication. The fact that the comp etito rs are consp ecifi cs cause s a ‘‘ ratc het effect’’ in which intelligence increases continually. Support for this line of theorizing has come from finding that the size of the primate brain—specifically, the neocortex—was closely related to measures of social group size (Dunbar, 1992a, 1993, 1995, 1998 ) and so ci al skil l ( Byr ne and Cor p, 2004; Rea der and Lal and , 2001; Reader and Lefebvre, 2001 ). Neuron 65, March 25, 2010 ª2010 Elsevier Inc. 815
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Primate Social Cognition:Uniquely Primate, Uniquely Social, or Just Unique?
Richard W. Byrne1,* and Lucy A. Bates1
1Centre for Social Learning and Cognitive Evolution and Scottish Primate Research Group, University of St Andrews, St Andrews KY16 9JP,Scotland, UK*Correspondence: [email protected] 10.1016/j.neuron.2010.03.010
Primates undoubtedly have impressive abilities in perceiving, recognizing, manipulating, and predicting
other individuals, but only great apes seem to recognize the cognitive basis of manipulative and cooperative
tactics or the concept of self. None of these abilities is unique to primates. We distinguish (1) a package of
quantitative advantages in social sophistication, perhaps based on more efficient memory, in which neocor-
tical enlargement is associated with the challenge of social living; from (2) a qualitative difference in under-
standing, whose taxonomic distribution—including several distantly related species, including birds—
does not point to an evolutionary origin in social challenges and may instead relate to a need to acquire novelways of dealing with the physical world. The ability of great apes to learn new manual routines by parsing
action components may have driven their qualitatively greater social skill, suggesting that strict partition
of physical and social cognition is likely to be misleading.
Introduction
Fifty years ago, human experimental psychology was dominated
by learning theory: but the ‘‘cognitive revolution’’ dramatically
changed the nature of research into human behavior. Since
then, neuroscientists have regularly used nonhuman primates
as a ‘‘simpler’’ model to test theories of human social cognition
and information processing, with most laboratory studies of
primates relying on a few species of macaque monkeys. During
the same period, a growing number of psychologists, anthropol-
ogists, and ethologists began studying the social behavior of
a wide range of nonhuman primates,both in theprimates’ natural
habitat and in the laboratory, in a parallel attempt to reconstruct
the evolutionary history of human mental competence.
In this review, we shall attempt to explain what these wider-
ranging studies have shown us about nonhuman primate social
cognition, in the hope that it will inform understanding in neuro-
science and new work on humans. It is important to note that
there are numerous differences in behavior between different
primate species, even among different species of monkeys—
as well as between monkeys and great apes, of which humans
are one of several extant species, including orangutans, gorillas,
and chimpanzees. Thesedifferences may reflect variations in the
cognitive architecture, or they may result from socioecologicaldifferences between species that limit the expression of similar
underlying cognitive capacity. Where appropriate, we therefore
identify the primate species studied, in order to avoid the trap
of treating all monkey species, or even all nonhuman primates,
as alike in their behavior and cognition.
Mimicking the paradigm shift in human experimental
psychology, there has been a later and more gradual change
toward cognitive explanations of animal behavior.Earlierdebates
about behavior were often dominated by the issue of whether
‘‘simple’’ associative learning could account for the data or
whether the animals ‘‘were cognitive,’’ usually taken to mean
having conscious thought processes ( Byrne and Bates, 2006;
Macphail, 1998 ). As in the case of human experimental
psychology, the associative approach was not refuted in prin-
ciple, but the sheer effort and increasingly ad hoc appearance
of associationist explanations for complex behavior led to the
ascendance of cognitive theorizing. A particular impetus for
change came from the growing evidence of sophistication and
complexity in primate behavior: in taking account of a rich
network of relationships ( Cheney and Seyfarth, 1982a, 1986,
1990 ), in interpreting others’ intentions ( Menzel, 1971, 1974;
Premack, 1988; Premack and Woodruff, 1978 ), and in deploying
subtle manipulative tactics ( Byrne and Whiten, 1985, 1991,
1992; de Waal, 1982, 1986; Whiten and Byrne, 1988 ). By the
mid-1990s, it had become acceptable to compare across
species by taking a generallycognitive approach, framed in terms
of information processing rather than consciousness ( Byrne,
1995; Tomasello and Call, 1997 ). This shift toward cognitive
explanations of behavior has now left us much better placed to
generate and test hypotheses about the evolution of human
cognitive skill.
Much of the new evidence of complexity in primate behavior
came from the social realm, and comparable sophistication
was not known for physical cognition ( Byrne and Whiten, 1988;
Cheney and Seyfarth, 1985; Jolly, 1966 ). This pattern meshedwith Humphrey’s influential theory ( Humphrey, 1976, 1981 ) that
the evolutionary challenge for which the solution is ‘‘intelligence’’
is more likely a social than a physical one. He argued that semi-
permanent social living, as found in many primate species, sets
up a selective pressure for increasing social sophistication. The
fact that the competitors are conspecifics causes a ‘‘ratchet
effect’’ in which intelligence increases continually. Support for
this line of theorizing has come from finding that the size of the
primate brain—specifically, the neocortex—was closely related
to measures of social group size ( Dunbar, 1992a, 1993, 1995,
1998 ) and social skill ( Byrne and Corp, 2004; Reader and Laland,
the failures and inadequacies become the stumbling block for
comparative explanation, rather than the successes. On current
evidence, only a few species show ‘‘extra,’’ qualitatively different
social capacities compared to most mammals and birds: why
just them? Dealing with this concern may require a more Kantianformulation, in which some crucial mental structure(s) must be
innate, and differ qualitatively between taxa, whereas statistical
extraction of regularities ‘‘fills in the gaps’’ from practical experi-
ence. In an analogy from the physical domain, consider
monkeys’ representation of causal properties. Tamarin monkeys
are sensitive to the properties of material that are relevant to
tool-using, such as rigidity and length, but not color and texture,
despite not naturally using tools ( Hauser, 1997 ). It seems that
monkeys are predisposed to represent properties that will—in
some species, in some environments—permit intelligent tool
use to be acquired. But in species that lack an innate predispo-
sition to represent the correct properties of materials, intelligent
tool use will never be developed.
What innate mental structure(s) might allow certain animaltaxa to develop ‘‘extra’’ capacities in the domain of social
behavior? In principle, ‘‘structure’’ can be provided in two inde-
pendent ways. The simpler way involves innate heuristics. For
instance, a well-known principle in animal learning is that innate
‘‘constraints’’ may focus learning on biologically useful things,
determining what is learned easily andwhat is not: rats associate
nauseabut not electric shock with novel foods eaten many hours
earlier ( Garcia et al., 1966 ); perhaps similar genetic channeling
determines that we so readily learn to fear spiders but have
to be strictly taught to fear electricity. At a more general level,
the degree of native curiosity and the tendency to latent learn-
ing, picking up useful knowledge from certain situations, will
be greater in species evolved to fill generalist niches in fast-
changing environments. Moreover, species that depend on
motor or social skills may be naturally playful during the juvenile
period, the play tending to allow development of an augmented
motor repertoire and social skills ( Bruner, 1972; Caro, 1995;
Fagen, 1976; Whiten and Byrne, 1991 ).
This level of explanation can account for some of the quanti-
tative differences between species, for instance those depen-
dent upon the complexity of social system and fineness of
social categorization. When it comes to species differences
that imply deeper levels of understanding, it may be necessary
to invoke differences in cognitive architecture. For instance,
social transmission of information occurs in monkeys and
great apes, but the effects are apparently very different, with
monkeys unable to learn complex novel behavior by imitation( Byrne, 2002a; Visalberghi and Fragaszy, 2000 ). Is this because
they have different cognitive systems? If species are not able
to form the same mental representations, then what they
can do with the same information may be radically different
( Byrne et al., 2004 ). For instance, any social vertebrate is likely
to benefit from social learning in some way, but those species
that code what they see in terms only of a ‘‘conspecific
engaged in activity at a place’’ may only gain from local
enhancement ( Hoppitt and Laland, 2008 ). In contrast, if a
species is able to code the time sequence of specific actions
applied by that conspecific, what it learns will be much richer
( Byrne, 2002b ).
We propose that the distribution of behavior-parsing abilities
will be critical to species differences, both directly and indirectly.
Direct benefits accrue when understanding of others’ actions is
required to deal with challenges: for instance, when correct
interpretation of a social situation requires working out the inten-tion behind another’s actions, or in imitatively acquiring a
complex skill that would be unlikely to be discovered by trial
and error exploration. In order to use the gist of another’s
behavior as a structure with which to build novel actions, it will
be essential to have, in tandem with the perceptual process of
behavior parsing, the ability to build up hierarchically organized
programs of goal-directed action, which will otherwise be
evident in general problem-solving behavior: planning capabil-
ities. Indirect benefits come from the informational content that
can be passed on by social learning and culture, allowing adap-
tation to local environments to be mediated by social learning as
well as genetical evolution ( van Schaik et al., 1999; Washburn
and Benedict, 1979; Whiten, 2000 ).
In summary, we suggest that social cognition is not uniqueto primates and that primate cognition is not uniquely social.
We argue that most demonstrations of cognitive skill can be
accounted for by quantitative differences in memory, with the
tendency to larger memory most likely being driven by social
competition (from conspecifics and from predator/prey interac-
tions). However, some particular skills such as insightful cooper-
ation or deception, perception of intent, imitation of novel skills,
and mirror self-recognition, signify a qualitatively different repre-
sentation of mechanisms and minds. This probably relies on the
presence of specific cognitive architecture that allows for
behavior parsing and the formation of hierarchically organized
programs of action, theselection for which mayhave been driven
by physical constraints. Our account of behavior parsing has
necessarily been focused narrowly on primates: the evidence
is just not there for other species, including those unusual
species that show convergent abilitieswith great apes. The chal-
lenge for futureresearchwill be to discover whether this explana-
tion might apply more generally or whether a wholly new theory
of higher-order cognitive abilities will need to be developed to
explain the full pattern.
ACKNOWLEDGMENTS
This paper has benefited greatly from the comments of three anonymous
referees, and we gratefully recognize their help; the errors and biases that
remain are wholly our own.
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