Alex Kacelnik

The Value of a Smile: Game Theory with a Human Face

Many economists and biologists view cooperation as anomalous: animals (including humans) that pursue their own self-interest have superior survival odds to their altruistic or cooperative neighbors. However, in many situations there are substantial gains to the group that can achieve cooperation among its members, and to individuals who are members of those groups. For an individual, the key to successful cooperation is the ability to identify cooperative partners. The ability to signal and detect the intention to cooperate would be a very valuable skill for humans to posses. Smiling is frequently observed in social interactions between humans, and may be used as a signal of the intention to cooperate. However, given that humans have the ability to smile falsely, the ability to detect intentions may go far beyond the ability to recognize a smile. In the present study, we examine the value of a smile in a simple bargaining context. 120 subjects participate in a laboratory experiment consisting of a simple two-person, one-shot “trust” game with monetary payoffs. Each subject is shown a photograph of his partner prior to the game; the photograph is taken from a collection that includes one smiling and one non-smiling image for each of 60 individuals. These photographs are also rated by a separate set of subjects who complete a semantic differential survey on affective and behavioral interpretations of the images. Results lend some support to the prediction that smiles can elicit cooperation among strangers in a one-shot interaction. Other characteristics of faces also appear to elicit cooperation. Factor analysis of the survey data reveals an important factor, termed “cooperation”, which is strongly related to trusting behavior in the game. This factor is correlated with smiling, but is somewhat more strongly predictive of behavior than a smile alone. In addition, males are found to be more cooperative, especially towards female images, whereas females are least cooperative towards female images.

Chick Begging Strategies in Relation to Brood Hierarchies and Hatching Asynchrony

Altricial offspring solicit food by begging, and their parents feed them according to begging intensity, which has been shown to be positively related to offspring need. Parent‐offspring genetic conflict calls for analyses of evolutionary stability, and various theoretical models have shown that stability is possible in the framework of handicap theory. The models predict that a negative relationship exists between offspring condition and begging and that offspring in poorer condition should be fed preferentially. However, these predictions depend on two unsatisfactory assumptions. First, they assume a monotonically decelerated relation between condition and fitness (this function is more likely to be sigmoid); second, they ignore physical competition between siblings, which is known to be important. We examined the significance of these issues by manipulating hatching asynchrony in broods of starlings Sturnus vulgaris, thus controlling competitive asymmetries between nest mates. We created broods with senior (older) and junior (younger) chicks and control broods with synchronous chicks. In field and laboratory experiments, we found that seniors begged less than juniors and controls, whereas juniors did not differ significantly from controls. However, seniors received more food from their parents and grew better than juniors or controls (hence, they were in better condition). These results violate the predictions of available theoretical models and, together with limitations in the universality of their assumptions, indicate that fundamental aspects of parent‐offspring communication are not yet understood.

Food allocation among nestling starlings : sibling competition and the scope of parental choice

Parental provisioning for nestling birds is generally considered to be an interactive, conflictive process because the optimal provisioning rate differs between parents and young and because nestlings are engaged in intersibling competition. Understanding the evolution of communication in such a situation presents unusual problems because the scope for parental strategies may be limited by competitive behaviour of the chicks. We substantiate this view by studying parent-offspring feeding interactions between chicks and provisioning adults in the European starling Sturnus vulgaris in relation to chick state and intersibling competition. The state of one target chick in each nest was manipulated in the field by temporarily placing it in enlarged, reduced or normal-sized broods before returning it to its original nest. Conditions in the original nests were standardized during manipulation by using substitute chicks. Once returned to its original brood, the probability of the target chick being fed increased if it increased its begging intensity and/or it positioned itself closer to the entrance of the nest. Both begging intensity and position were functions of the treatment previously experienced, with target chicks begging more and attaining positions closer to the nest entrance after they had spent time in larger broods. We postulate that these factors must be included in theoretical analyses of the evolution of food-solicitation signalling because, although the effect of begging on feeding probability may be mediated by parental choice, the effect of position depends on between-chick dynamics, and the parents apparently accept the outcome of these interactions.

Cost can increase preference in starlings

We used European starlings, Sturnus vulgaris, to investigate the relationship between the cost paid to obtain food rewards and preference between stimuli associated with the resulting rewards. In no-choice trials either 16 1-m flights (high effort) or four 1-m flights (low effort) gave access to differently coloured keys. Pecking at these keys resulted in identical food rewards. When subjects were given choices between the coloured keys in choice trials without having paid any effort, the majority preferred the coloured key that was paired with the higher level of work in no-choice trials. We relate our findings to results in animal behaviour, psychology and economics, and give a theoretical account that has implications for phenomena variously recognized as the ‘sunk cost fallacy’ (the tendency to invest more in something after much has already been invested), ‘work ethics’ (valuing an option more as a result of physical effort), ‘cognitive dissonance’ (making mental effort to overlook or re-evaluate information that does not accord with a dominant internal representation) and the ‘Concorde Fallacy’ (the readiness to forego more fitness for something that has been responsible for greater fitness compromise in the past).

Development of tool use in New Caledonian crows: inherited action patterns and social influences

New Caledonian crows, Corvus moneduloides, are the most advanced avian tool makers and tool users. We previously reported that captive-bred isolated New Caledonian crows spontaneously use twig tools and cut tools out of Pandanus spp. tree leaves, an activity possibly under cultural influence in the wild. However, what aspects of these behaviours are inherited and how they interact with individual and social experience remained unknown. To examine the interaction between inherited traits, individual learning and social transmission, we observed the ontogeny of twig tool use in hand-reared juveniles. Successful food retrieval was preceded by stereotyped object manipulation action patterns that resembled components of the mature behaviour, demonstrating that tool-oriented behaviours in this species are an evolved specialization. However, there was also an effect of social learning: juveniles that had received demonstrations of twig tool use by their human foster parent showed higher levels of handling and insertion of twigs than did their naive counterparts; a choice experiment showed that they preferred to handle objects that they had seen being manipulated by their human foster parent. Our observations are consistent with the hypothesis that individual learning, cultural transmission and creative problem solving all contribute to the acquisition of the tool-oriented behaviours in the wild, but inherited species-typical action patterns have a greater role than has been recognized.

Framing effects and risky decisions in starlings

Animals are predominantly risk prone toward reward delays and risk averse toward reward amounts. Humans in turn tend to be risk-seeking for losses and risk averse for gains. To explain the human results, Prospect Theory postulates a convex utility for losses and concave utility for gains. In contrast, Scalar Utility Theory (SUT) explains the animal data by postulating that the cognitive representation of outcomes follows Webers Law, namely that the spread of the distribution of expected outcomes is proportional to its mean. SUT also would explain human results if utility (even if it is linear on expected outcome) followed Webers Law. We present an experiment that simulates losses and gains in a bird, the European Starling, to test the implication of SUT that risk proneness/aversion should extend to any aversive/desirable dimension other than time and amount of reward. Losses and gains were simulated by offering choices of fixed vs. variable outcomes with lower or higher outcomes than what the birds expected. The subjects were significantly more risk prone for losses than for gains but, against expectations, they were not significantly risk averse toward gains. The results are thus, in part, consistent with Prospect Theory and SUT and show that risk attitude in humans and birds may obey a common fundamental principle.

Visual attention and the acquisition of information in human crowds

Pedestrian crowds can form the substrate of important socially contagious behaviors, including propagation of visual attention, violence, opinions, and emotional state. However, relating individual to collective behavior is often difficult, and quantitative studies have largely used laboratory experimentation. We present two studies in which we tracked the motion and head direction of 3,325 pedestrians in natural crowds to quantify the extent, influence, and context dependence of socially transmitted visual attention. In our first study, we instructed stimulus groups of confederates within a crowd to gaze up to a single point atop of a building. Analysis of passersby shows that visual attention spreads unevenly in space and that the probability of pedestrians adopting this behavior increases as a function of stimulus group size before saturating for larger groups. We develop a model that predicts that this gaze response will lead to the transfer of visual attention between crowd members, but it is not sufficiently strong to produce a tipping point or critical mass of gaze-following that has previously been predicted for crowd dynamics. A second experiment, in which passersby were presented with two stimulus confederates performing suspicious/irregular activity, supports the predictions of our model. This experiment reveals that visual interactions between pedestrians occur primarily within a 2-m range and that gaze-copying, although relatively weak, can facilitate response to relevant stimuli. Although the above aspects of gaze-following response are reproduced robustly between experimental setups, the overall tendency to respond to a stimulus is dependent on spatial features, social context, and sex of the passerby.

State-dependent learned valuation drives choice in an invertebrate

Humans and other vertebrates occasionally show a preference for items remembered to be costly or experienced when the subject was in a poor condition (this is known as a sunk-costs fallacy or state-dependent valuation). Whether these mechanisms shared across vertebrates are the result of convergence toward an adaptive solution or evolutionary relicts reflecting common ancestral traits is unknown. Here we show that state-dependent valuation also occurs in an invertebrate, the desert locust Schistocerca gregaria (Orthoptera: Acrididae). Given the latters phylogenetic and neurobiological distance from those groups in which the phenomenon was already known, we suggest that state-dependent valuation mechanisms are probably ecologically rational solutions to widespread problems of choice.

Selection of tool diameter by New Caledonian crows Corvus moneduloides

One important element of complex and flexible tool use, particularly where tool manufacture is involved, is the ability to select or manufacture appropriate tools anticipating the needs of any given task—an ability that has been rarely tested in non-primates. We examine aspects of this ability in New Caledonian crows—a species known to be extraordinary tool users and manufacturers. In a 2002 study, Chappell and Kacelnik showed that these crows were able to select a tool of the appropriate length for a task among a set of different lengths, and in 2002, Weir, Chappell and Kacelnik showed that New Caledonian crows were able to shape unfamiliar materials to create a usable tool for a specific task. Here we examine their handling of tool diameter. In experiment 1, we show that when facing three loose sticks that were usable as tools, they preferred the thinnest one. When the three sticks were presented so that one was loose and the other two in a bundle, they only disassembled the bundle when their preferred tool was tied. In experiment 2, we show that they manufacture, and modify during use, a tool of a suitable diameter from a tree branch, according to the diameter of the hole through which the tool will have to be inserted. These results add to the developing picture of New Caledonian crows as sophisticated tool users and manufacturers, having an advanced level of folk physics.

Central-place foraging in honey bees: the effect of travel time and nectar flow on crop filling

SummaryCrop-filling by honeybees foraging at sources of variable nectar flow at a fixed distance from the hive has been shown to maximize energetic efficiency, defined as ratio of energy gained to energy spent. Predictions based on maximisation of rate of energy gain, defined as net energy gained per unit time foraging, are significantly different from observed behaviour (Schmid-Hempel et al. 1985). In this paper we consider the effect of varying travel times in addition to flow rate. The predictions of an extended version of our theoretical model are confronted with experimental results obtained by Núñez (1982). Núñez found that bees filled their crops more fully for higher flows and longer travel times. We show that when the cost of carrying a load is considered, this trend can be predicted by maximising either energetic efficiency or net rate of gain. Figure 1 shows, however, that maximisation of net rate of gain can only produce an acceptable quantitative fit if unreasonably high costs are assumed to result from carrying the load. Energetic efficiency instead generates a good quantitative fit for acceptable assumptions about this cost (Fig. 2).