Laurie R. Santos

How Basic Are Behavioral Biases? Evidence from Capuchin Monkey Trading Behavior

Behavioral economics has demonstrated systematic decision‐making biases in both lab and field data. Do these biases extend across contexts, cultures, or even species? We investigate this question by introducing fiat currency and trade to a colony of capuchin monkeys and recovering their preferences over a range of goods and gambles. We show that capuchins react rationally to both price and wealth shocks but display several hallmark biases when faced with gambles, including reference dependence and loss aversion. Given our capuchins’ inexperience with money and trade, these results suggest that loss aversion extends beyond humans and may be innate rather than learned.

The Evolution of Self-Control

Significance Although scientists have identified surprising cognitive flexibility in animals and potentially unique features of human psychology, we know less about the selective forces that favor cognitive evolution, or the proximate biological mechanisms underlying this process. We tested 36 species in two problem-solving tasks measuring self-control and evaluated the leading hypotheses regarding how and why cognition evolves. Across species, differences in absolute (not relative) brain volume best predicted performance on these tasks. Within primates, dietary breadth also predicted cognitive performance, whereas social group size did not. These results suggest that increases in absolute brain size provided the biological foundation for evolutionary increases in self-control, and implicate species differences in feeding ecology as a potential selective pressure favoring these skills. Cognition presents evolutionary research with one of its greatest challenges. Cognitive evolution has been explained at the proximate level by shifts in absolute and relative brain volume and at the ultimate level by differences in social and dietary complexity. However, no study has integrated the experimental and phylogenetic approach at the scale required to rigorously test these explanations. Instead, previous research has largely relied on various measures of brain size as proxies for cognitive abilities. We experimentally evaluated these major evolutionary explanations by quantitatively comparing the cognitive performance of 567 individuals representing 36 species on two problem-solving tasks measuring self-control. Phylogenetic analysis revealed that absolute brain volume best predicted performance across species and accounted for considerably more variance than brain volume controlling for body mass. This result corroborates recent advances in evolutionary neurobiology and illustrates the cognitive consequences of cortical reorganization through increases in brain volume. Within primates, dietary breadth but not social group size was a strong predictor of species differences in self-control. Our results implicate robust evolutionary relationships between dietary breadth, absolute brain volume, and self-control. These findings provide a significant first step toward quantifying the primate cognitive phenome and explaining the process of cognitive evolution.

Primate brains in the wild: the sensory bases for social interactions.

Each organism in the animal kingdom has evolved to detect and process a specific set of stimuli in its environment. Studies of an animals socioecology can help us to identify these stimuli, as well as the natural behavioural responses that they evoke and control. Primates are no exception, but many of our specializations are in the social domain. How did the human brain come to be so exquisitely tuned to social interactions? Only a comparative approach will provide the answer. Behavioural studies are shedding light on the sensory bases for non-human primate social interactions, and data from these studies are paving the way for investigations into the neural bases of sociality.

Endowment effect in capuchin monkeys.

In humans, the capacity for economically rational choice is constrained by a variety of preference biases: humans evaluate gambles relative to arbitrary reference points; weigh losses heavier than equally sized gains; and demand a higher price for owned goods than for equally preferred goods that are not yet owned. To date, however, fewer studies have examined the origins of these biases. Here, we review previous work demonstrating that human economic biases such as loss aversion and reference dependence are shared with an ancestrally related New World primate, the capuchin monkey (Cebus apella). We then examine whether capuchins display an endowment effect in a token-trading task. We identified pairs of treats (fruit discs versus cereal chunks) that were equally preferred by each monkey. When given a chance to trade away their owned fruit discs to obtain the equally valued cereal chunks (or vice versa), however, monkeys required a far greater compensation than the equally preferred treat. We show that these effects are not due to transaction costs or timing issues. These data suggest that biased preferences rely on cognitive systems that are more evolutionarily ancient than previously thought—and that common evolutionary ancestry shared by humans and capuchins may account for the occurrence of the endowment effect in both species.

Young Children Are More Generous When Others Are Aware of Their Actions

Adults frequently employ reputation-enhancing strategies when engaging in prosocial acts, behaving more generously when their actions are likely to be witnessed by others and even more so when the extent of their generosity is made public. This study examined the developmental origins of sensitivity to cues associated with reputationally motivated prosociality by presenting five-year-olds with the option to provide one or four stickers to a familiar peer recipient at no cost to themselves. We systematically manipulated the recipient’s knowledge of the actor’s choices in two different ways: (1) occluding the recipient’s view of both the actor and the allocation options and (2) presenting allocations in opaque containers whose contents were visible only to the actor. Children were consistently generous only when the recipient was fully aware of the donation options; in all cases in which the recipient was not aware of the donation options, children were strikingly ungenerous. These results demonstrate that five-year-olds exhibit “strategic prosociality,” behaving differentially generous as a function of the amount of information available to the recipient about their actions. These findings suggest that long before they develop a rich understanding of the social significance of reputation or are conscious of complex strategic reasoning, children behave more generously when the details of their prosocial actions are available to others.

The evolution of intergroup bias: Perceptions and attitudes in rhesus macaques.

Social psychologists have learned a great deal about the nature of intergroup conflict and the attitudinal and cognitive processes that enable it. Less is known about where these processes come from in the first place. In particular, do our strategies for dealing with other groups emerge in the absence of human-specific experiences? One profitable way to answer this question has involved administering tests that are conceptual equivalents of those used with adult humans in other species, thereby exploring the continuity or discontinuity of psychological processes. We examined intergroup preferences in a nonhuman species, the rhesus macaque (Macaca mulatta). We found the first evidence that a nonhuman species automatically distinguishes the faces of members of its own social group from those in other groups and displays greater vigilance toward outgroup members (Experiments 1-3). In addition, we observed that macaques spontaneously associate novel objects with specific social groups and display greater vigilance to objects associated with outgroup members (Experiments 4-5). Finally, we developed a looking time procedure-the Looking Time Implicit Association Test, which resembles the Implicit Association Test (Greenwald & Banaji, 1995)-and we discovered that macaques, like humans, automatically evaluate ingroup members positively and outgroup members negatively (Experiments 6-7). These field studies represent the first controlled experiments to examine the presence of intergroup attitudes in a nonhuman species. As such, these studies suggest that the architecture of the mind that enables the formation of these biases may be rooted in phylogenetically ancient mechanisms.

Object individuation using property/kind information in rhesus macaques (Macaca mulatta)

Around 1 year of age, infants develop the ability to individuate objects in the absence of spatiotemporal information. Some have proposed that this capacity relies on the emergence of language and, in particular, that comprehending an objects label is required to individuate it as a particular kind. One approach to testing this hypothesis is to conduct experiments on pre-linguistic human infants. A second is to test non-linguistic animals. We followed the second approach, exploring whether semi-free-ranging rhesus macaques can individuate objects using property/kind information. To make the results most directly comparable, we adapted a reaching paradigm used to examine property/kind individuation in infants. Results from three experiments demonstrate that, like 12-month-old infants, adult rhesus macaques can use both spatiotemporal and property/kind information to individuate food objects. In a fourth experiment designed to examine which properties are used to individuate food objects, results revealed that rhesus use color, but not shape. These results, together with experiments involving different procedures, provide support for the conclusion that in the absence of linguistic abilities, some non-human primates spontaneously use property/kind information to individuate objects.

Expectations about numerical events in four lemur species (Eulemur fulvus, Eulemur mongoz, Lemur catta and Varecia rubra)

Although much is known about how some primates—in particular, monkeys and apes—represent and enumerate different numbers of objects, very little is known about the numerical abilities of prosimian primates. Here, we explore how four lemur species (Eulemur fulvus, E. mongoz, Lemur catta, and Varecia rubra) represent small numbers of objects. Specifically, we presented lemurs with three expectancy violation looking time experiments aimed at exploring their expectations about a simple 1+1 addition event. In these experiments, we presented subjects with displays in which two lemons were sequentially added behind an occluder and then measured subjects’ duration of looking to expected and unexpected outcomes. In experiment 1, subjects looked reliably longer at an unexpected outcome of only one object than at an expected outcome of two objects. Similarly, subjects in experiment 2 looked reliably longer at an unexpected outcome of three objects than at an expected outcome of two objects. In experiment 3, subjects looked reliably longer at an unexpected outcome of one object twice the size of the original than at an expected outcome of two objects of the original size. These results suggest that some prosimian primates understand the outcome of simple arithmetic operations. These results are discussed in light of similar findings in human infants and other adult primates.

How monkeys see the eyes: cotton-top tamarins' reaction to changes in visual attention and action

Among social species, the capacity to detect where another individual is looking is adaptive because gaze direction often predicts what an individual is attending to, and thus what its future actions are likely to be. We used an expectancy violation procedure to determine whether cotton-top tamarins (Saguinus oedipus oedipus) use the direction of another individual’s gaze to predict future actions. Subjects were familiarized with a sequence in which a human actor turned her attention toward one of two objects sitting on a table and then reached for that object. Following familiarization, subjects saw two test events. In one test event, the actor gazed at the new object and then reached for that object. From a human perspective, this event is considered consistent with the causal relationship between visual attention and subsequent action, that is, grabbing the object attended to. In the second test event, the actor gazed at the old object, but reached for the new object. This event is considered a violation of expectation. When the actor oriented with both her head-and-eyes, subjects looked significantly longer at the second test event in which the actor reached for the object to which she had not previously oriented. However, there was no difference in looking time between test events when the actor used only her eyes to orient. These findings suggest that tamarins are able to use some combination of head orientation and gaze direction, but not gaze direction alone, to predict the actions of a human agent.

Probing the limits of tool competence: Experiments with two non-tool-using species (Cercopithecus aethiops and Saguinus oedipus)

Non-human animals vary in their ability to make and use tools. The goal of the present study was to further explore what, if anything, differs between tool-users and non-tool-users, and whether these differences lie in the conceptual or motor domain. We tested two species that typically do not use tools—cotton top tamarins (Saguinus oedipus) and vervet monkeys (Cercopithecus aethiops)—on problems that mirrored those designed for prolific tool users such as chimpanzees. We trained subjects on a task in which they could choose one of two canes to obtain an out-of-reach food reward. After training, subjects received several variations on the original task, each designed to examine a specific conceptual aspect of the pulling problem previously studied in other tool-using species. Both species recognized that effective pulling tools must be made of rigid materials. Subsequent conditions revealed significant species differences, with vervets outperforming tamarins across many conditions. Vervets, but not tamarins, had some recognition of the relationship between a tools orientation and the position of the food reward, the relationship between a tools trajectory and the substance that it moves on, and that tools must be connected in order to work properly. These results provide further evidence that tool-use may derive from domain-general, rather than domain-specific cognitive capacities that evolved for tool use per se.