Monkeys Are More Patient in a Foraging Task than in a Standard Intertemporal Choice Task

Studies of animal impulsivity generally find steep subjective devaluation, or discounting, of delayed rewards – often on the order of a 50% reduction in value in a few seconds. Because such steep discounting is highly disfavored in evolutionary models of time preference, we hypothesize that discounting tasks provide a poor measure of animals’ true time preferences. One prediction of this hypothesis is that estimates of time preferences based on these tasks will lack external validity, i.e. fail to predict time preferences in other contexts. We examined choices made by four rhesus monkeys in a computerized patch-leaving foraging task interleaved with a standard intertemporal choice task. Monkeys were significantly more patient in the foraging task than in the intertemporal choice task. Patch-leaving behavior was well fit by parameter-free optimal foraging equations but poorly fit by the hyperbolic discount parameter obtained from the intertemporal choice task. Day-to-day variation in time preferences across the two tasks was uncorrelated with each other. These data are consistent with the conjecture that seemingly impulsive behavior in animals is an artifact of their difficulty understanding the structure of intertemporal choice tasks, and support the idea that animals are more efficient rate maximizers in the multi-second range than intertemporal choice tasks would suggest.     

Evolution of Self-Organized Task Specialization in Robot Swarms

Division of labor is ubiquitous in biological systems, as evidenced by various forms of complex task specialization observed in both animal societies and multicellular organisms. Although clearly adaptive, the way in which division of labor first evolved remains enigmatic, as it requires the simultaneous co-occurrence of several complex traits to achieve the required degree of coordination. Recently, evolutionary swarm robotics has emerged as an excellent test bed to study the evolution of coordinated group-level behavior. Here we use this framework for the first time to study the evolutionary origin of behavioral task specialization among groups of identical robots. The scenario we study involves an advanced form of division of labor, common in insect societies and known as “task partitioning”, whereby two sets of tasks have to be carried out in sequence by different individuals. Our results show that task partitioning is favored whenever the environment has features that, when exploited, reduce switching costs and increase the net efficiency of the group, and that an optimal mix of task specialists is achieved most readily when the behavioral repertoires aimed at carrying out the different subtasks are available as pre-adapted building blocks. Nevertheless, we also show for the first time that self-organized task specialization could be evolved entirely from scratch, starting only from basic, low-level behavioral primitives, using a nature-inspired evolutionary method known as Grammatical Evolution. Remarkably, division of labor was achieved merely by selecting on overall group performance, and without providing any prior information on how the global object retrieval task was best divided into smaller subtasks. We discuss the potential of our method for engineering adaptively behaving robot swarms and interpret our results in relation to the likely path that nature took to evolve complex sociality and task specialization.     

Impulsive Social Influence Increases Impulsive Choices on a Temporal Discounting Task in Young Adults

Adolescents and young adults who affiliate with friends who engage in impulsive behavior are more likely to engage in impulsive behaviors themselves, and those who associate with prosocial (i.e. more prudent, future oriented) peers are more likely to engage in prosocial behavior. However, it is difficult to disentangle the contribution of peer influence vs. peer selection (i.e., whether individuals choose friends with similar traits) when interpreting social behaviors. In this study, we combined a novel social manipulation with a well-validated delay discounting task assessing impulsive behavior to create a social influence delay discounting task, in which participants were exposed to both impulsive (smaller, sooner or SS payment) and non-impulsive (larger, later or LL payment) choices from their peers. Young adults in this sample, n = 51, aged 18–25 had a higher rate of SS choices after exposure to impulsive peer influence than after exposure to non-impulsive peer influence. Interestingly, in highly susceptible individuals, the rate of non-impulsive choices did not increase after exposure to non-impulsive influence. There was a positive correlation between self-reported suggestibility and degree of peer influence on SS choices. These results suggest that, in young adults, SS choices appear to be influenced by the choices of same-aged peers, especially for individuals who are highly susceptible to influence.     

Foraging for Work and Age-Based Polyethism: The Roles of Age and Previous Experience on Task Choice in Ants

In social insects, colonies commonly show temporal polyethism in worker behavior, such that a worker follows a predictable pattern of changes between tasks as it ages. This pattern usually leads from workers first doing a safe task like brood care, to ending their lives doing the most dangerous tasks like foraging. Two mechanisms could potentially underlie this pattern: (1) age‐based task allocation, where the aging process itself predisposes workers to switch to more dangerous tasks; and (2) foraging for work, where ants switch to tasks that need doing from tasks which have too many associated workers. We tested the relative influence of these mechanisms by establishing nests of Camponotus floridanus with predetermined combinations of workers of known age and previous task specialization. The results supported both mechanisms. Nests composed of entirely brood‐tending workers had the oldest workers preferentially switching to foraging. However, in nests initially composed entirely of foragers, the final distribution of tenders and foragers was not different from random task‐switching and therefore supportive of foraging for work. Thus, it appears that in C. floridanus there is directionality to the mechanisms of task allocation. Switching to more dangerous tasks is age‐influenced, but switching to less dangerous tasks is age‐independent. The results also suggest that older workers are more flexible in their task choice behavior. Younger workers are more biased towards choosing within‐nest tasks. Finally, there are effects of previous experience that tend to keep ants in familiar tasks. Task allocation based on several mechanisms may balance between: (1) concentrating the most worn workers into the most dangerous tasks; (2) increasing task performance levels; and (3) maintaining behavioral flexibility to respond to demographic perturbations. The degree to which behavior is flexible may correlate to the frequency of such perturbations in a species.     

A Leaky-Integrator Model as a Control Mechanism Underlying Flexible Decision Making during Task Switching

The ability to switch between tasks is critical for animals to behave according to context. Although the association between the prefrontal cortex and task switching has been well documented, the ultimate modulation of sensory–motor associations has yet to be determined. Here, we modeled the results of a previous study showing that task switching can be accomplished by communication from distinct populations of sensory neurons. We proposed a leaky-integrator model where relevant and irrelevant information were stored separately in two integrators and task switching was achieved by leaking information from the irrelevant integrator. The model successfully explained both the behavioral and neuronal data. Additionally, the leaky-integrator model showed better performance than an alternative model, where irrelevant information was discarded by decreasing the weight on irrelevant information, when animals initially failed to commit to a task. Overall, we propose that flexible switching is, in part, achieved by actively controlling the amount of leak of relevant and irrelevant information.     

Reproductive numbers for nonautonomous spatially distributed periodic SIS models acting on two time scales

In this work we deal with a general class of spatially distributed periodic SIS epidemic models with two time scales. We let susceptible and infected individuals migrate between patches with periodic time dependent migration rates. The existence of two time scales in the system allows to describe certain features of the asymptotic behavior of its solutions with the help of a less dimensional, aggregated, system. We derive global reproduction numbers governing the general spatially distributed nonautonomous system through the aggregated system. We apply this result when the mass action law and the frequency dependent transmission law are considered. Comparing these global reproductive numbers to their non spatially distributed counterparts yields the following: adequate periodic migration rates allow global persistence or eradication of epidemics where locally, in absence of migrations, the contrary is expected.     

An Experimental Study of Team Size and Performance on a Complex Task

The relationship between team size and productivity is a question of broad relevance across economics, psychology, and management science. For complex tasks, however, where both the potential benefits and costs of coordinated work increase with the number of workers, neither theoretical arguments nor empirical evidence consistently favor larger vs. smaller teams. Experimental findings, meanwhile, have relied on small groups and highly stylized tasks, hence are hard to generalize to realistic settings. Here we narrow the gap between real-world task complexity and experimental control, reporting results from an online experiment in which 47 teams of size ranging from n = 1 to 32 collaborated on a realistic crisis mapping task. We find that individuals in teams exerted lower overall effort than independent workers, in part by allocating their effort to less demanding (and less productive) sub-tasks; however, we also find that individuals in teams collaborated more with increasing team size. Directly comparing these competing effects, we find that the largest teams outperformed an equivalent number of independent workers, suggesting that gains to collaboration dominated losses to effort. Importantly, these teams also performed comparably to a field deployment of crisis mappers, suggesting that experiments of the type described here can help solve practical problems as well as advancing the science of collective intelligence.     

Percent lipid is associated with body size but not task in the bumble bee <Emphasis Type="Italic">Bombus impatiens</Emphasis>

In some group-living organisms, labor is divided among individuals. This allocation to particular tasks is frequently stable and predicted by individual physiology. Social insects are excellent model organisms in which to investigate the interplay between physiology and individual behavior, as division of labor is an important feature within colonies, and individual physiology varies among the highly related individuals of the colony. Previous studies have investigated what factors are important in determining how likely an individual is, compared to nestmates, to perform certain tasks. One such task is foraging. Corpulence (i.e., percent lipid) has been shown to determine foraging propensity in honey bees and ants, with leaner individuals being more likely to be foragers. Is this a general trend across all social insects? Here we report data analyzing the individual physiology, specifically the percent lipid, of worker bumble bees (Bombus impatiens) from whom we also analyze behavioral task data. Bumble bees are also unusual among the social bees in that workers may vary widely in size. Surprisingly we find that, unlike other social insects, percent lipid is not associated with task propensity. Rather, body size closely predicts individual relative lipid stores, with smaller worker bees being allometrically fatter than larger worker bees.     

Theory of Choice in Bandit,Information Sampling and Foraging Tasks

Decision making has been studied with a wide array of tasks. Here we examine the theoretical structure of bandit, information sampling and foraging tasks. These tasks move beyond tasks where the choice in the current trial does not affect future expected rewards. We have modeled these tasks using Markov decision processes (MDPs). MDPs provide a general framework for modeling tasks in which decisions affect the information on which future choices will be made. Under the assumption that agents are maximizing expected rewards, MDPs provide normative solutions. We find that all three classes of tasks pose choices among actions which trade-off immediate and future expected rewards. The tasks drive these trade-offs in unique ways, however. For bandit and information sampling tasks, increasing uncertainty or the time horizon shifts value to actions that pay-off in the future. Correspondingly, decreasing uncertainty increases the relative value of actions that pay-off immediately. For foraging tasks the time-horizon plays the dominant role, as choices do not affect future uncertainty in these tasks.     

Task Design Influences Prosociality in Captive Chimpanzees (Pan troglodytes)

Chimpanzees confer benefits on group members, both in the wild and in captive populations. Experimental studies of how animals allocate resources can provide useful insights about the motivations underlying prosocial behavior, and understanding the relationship between task design and prosocial behavior provides an important foundation for future research exploring these animals'' social preferences. A number of studies have been designed to assess chimpanzees'' preferences for outcomes that benefit others (prosocial preferences), but these studies vary greatly in both the results obtained and the methods used, and in most cases employ procedures that reduce critical features of naturalistic social interactions, such as partner choice. The focus of the current study is on understanding the link between experimental methodology and prosocial behavior in captive chimpanzees, rather than on describing these animals'' social motivations themselves. We introduce a task design that avoids isolating subjects and allows them to freely decide whether to participate in the experiment. We explore key elements of the methods utilized in previous experiments in an effort to evaluate two possibilities that have been offered to explain why different experimental designs produce different results: (a) chimpanzees are less likely to deliver food to others when they obtain food for themselves, and (b) evidence of prosociality may be obscured by more “complex” experimental apparatuses (e.g., those including more components or alternative choices). Our results suggest that the complexity of laboratory tasks may generate observed variation in prosocial behavior in laboratory experiments, and highlights the need for more naturalistic research designs while also providing one example of such a paradigm.     

Task partitioning in swarms of robots: an adaptive method for strategy selection

Task partitioning is the decomposition of a task into two or more sub-tasks that can be tackled separately. Task partitioning can be observed in many species of social insects, as it is often an advantageous way of organizing the work of a group of individuals. Potential advantages of task partitioning are, among others: reduction of interference between workers, exploitation of individuals?? skills and specializations, energy efficiency, and higher parallelism. Even though swarms of robots can benefit from task partitioning in the same way as social insects do, only few works in swarm robotics are dedicated to this subject. In this paper, we study the case in which a swarm of robots has to tackle a task that can be partitioned into a sequence of two sub-tasks. We propose a method that allows the individual robots in the swarm to decide whether to partition the given task or not. The method is self-organized, relies on the experience of each individual, and does not require explicit communication between robots. We evaluate the method in simulation experiments, using foraging as testbed. We study cases in which task partitioning is preferable and cases in which it is not. We show that the proposed method leads to good performance of the swarm in both cases, by employing task partitioning only when it is advantageous. We also show that the swarm is able to react to changes in the environmental conditions by adapting the behavior on-line. Scalability experiments show that the proposed method performs well across all the tested group sizes.     

Behavioural Lateralization in Budgerigars Varies with the Task and the Individual

Handedness/footedness and side biases are a well-known phenomenon in many animals, including humans. However, these so-called biases have mostly been studied at the population level - individual biases have received less attention, especially with regard to consistency over different tasks. Here we investigate behavioral lateralization in 12 male Budgerigars, Melopsittacus undulatus, a social parrot inhabiting the Australian bushlands. We performed 5 types of experiments to investigate lateralization, in tasks that involved climbing onto a perch, or landing on perches arranged in various configurations. The birds displayed highly significant, individually varying biases. The bias displayed by any particular individual varied with the task, in strength as well as polarity. Analysis of the data revealed that the preferred foot used for climbing did not coincide with the foot that was used while landing. Thus, landing choices are probably not determined by foot bias. Furthermore, these individual preferences were overridden completely when a bird had to perform a task simultaneously with another bird.     

Local orientation and the evolution of foraging: changes in decision making can eliminate evolutionary trade-offs

Information processing is a major aspect of the evolution of animal behavior. In foraging, responsiveness to local feeding opportunities can generate patterns of behavior which reflect or "recognize patterns" in the environment beyond the perception of individuals. Theory on the evolution of behavior generally neglects such opportunity-based adaptation. Using a spatial individual-based model we study the role of opportunity-based adaptation in the evolution of foraging, and how it depends on local decision making. We compare two model variants which differ in the individual decision making that can evolve (restricted and extended model), and study the evolution of simple foraging behavior in environments where food is distributed either uniformly or in patches. We find that opportunity-based adaptation and the pattern recognition it generates, plays an important role in foraging success, particularly in patchy environments where one of the main challenges is "staying in patches". In the restricted model this is achieved by genetic adaptation of move and search behavior, in light of a trade-off on within- and between-patch behavior. In the extended model this trade-off does not arise because decision making capabilities allow for differentiated behavioral patterns. As a consequence, it becomes possible for properties of movement to be specialized for detection of patches with more food, a larger scale information processing not present in the restricted model. Our results show that changes in decision making abilities can alter what kinds of pattern recognition are possible, eliminate an evolutionary trade-off and change the adaptive landscape.     

Applying evolutionary models to the laboratory study of social learning

Cultural evolution is driven, in part, by the strategies that individuals employ to acquire behavior from others. These strategies themselves are partly products of natural selection, making the study of social learning an inherently Darwinian project. Formal models of the evolution of social learning suggest that reliance on social learning should increase with task difficulty and decrease with the probability of environmental change. These models also make predictions about how individuals integrate information from multiple peers. We present the results of microsociety experiments designed to evaluate these predictions. The first experiment measures baseline individual learning strategy in a two-armed bandit environment with variation in task difficulty and temporal fluctuation in the payoffs of the options. Our second experiment addresses how people in the same environment use minimal social information from a single peer. Our third experiment expands on the second by allowing access to the behavior of several other individuals, permitting frequency-dependent strategies like conformity. In each of these experiments, we vary task difficulty and environmental fluctuation. We present several candidate strategies and compute the expected payoffs to each in our experimental environment. We then fit to the data the different models of the use of social information and identify the best-fitting model via model comparison techniques. We find substantial evidence of both conformist and nonconformist social learning and compare our results to theoretical expectations.     

The role of trade-off shapes in the evolution of parasites in spatial host populations: an approximate analytical approach

Given the substantial changes in mixing in many populations, there is considerable interest in the role that spatial structure can play in the evolution of disease. Here we examine the role of different trade-off shapes in the evolution of parasites in a spatially structured host population where infection can occur locally or globally. We develop an approximate adaptive dynamic analytical approach, to examine how the evolutionarily stable (ES) virulence depends not only on the fraction of global infection/transmission but also on the shape of the trade-off between transmission and virulence. Our analysis can successfully predict the ES virulence found previously by simulation of the full system. The analysis confirms that when there is a linear trade-off between transmission and virulence spatial structure may lead to an ES virulence that increases as the proportion of global transmission increases. However, we also show that the ESS disappears above a threshold level of global infection, leading to maximization. In addition just below this threshold, there is the possibility of evolutionary bi-stabilities. When we assume the realistic trade-off between transmission and virulence that results in an ESS in the classical mixed model, we find that spatial structure can increase or decrease the ES virulence. A relatively high proportion of local infection reduces virulence but intermediate levels can select for higher virulence. Our work not only emphasizes the importance of spatial structure to the evolution of parasites, but also makes it clear that situations between the local and the global need to be considered. We also emphasize the key role that the shape of trade-offs plays in evolutionary outcomes.     

Social dominance, task performance and nutrition: implications for reproduction in eusocial wasps

Dominance status is associated with individual differences in reproductive capacity in many animal societies, but the mechanisms that link social dominance to reproductive physiology are poorly understood. We propose a model for social dynamics that incorporates the nutritional costs and benefits of behavior: dominant individuals avoid energy-expensive behavior and build their nutritional reserves, thereby increasing their potential for reproduction. Greater reproductive capacity, once achieved, favors increased social dominance. To test the model, we measured relationships of females' nutrient storage and reproductive capacities with dominance status and task performance in the eusocial wasp Mischocyttarus mastigophorus. Ovary development was positively related with high levels of nutrient storage and with high rates of dominance behavior, but was not correlated with task performance. In contrast, high levels of nutrient storage were positively related with the performance of nutrient garnering and conserving tasks, but not with dominance behavior. These data support a model which places the nutritional costs of task performance as an intermediate causal link that connects dominance status with the accumulation of nutrient stores. Nutrient flow may be a general causal mechanism linking dominance status to reproductive capacity in animal societies.     

Approximate aggregation of a two time scales periodic multi-strain SIS epidemic model: A patchy environment with fast migrations

In this work we consider a spatially distributed periodic multi strain SIS epidemic model. We let susceptible and infected individuals migrate between patches, with periodic migration rates. Considering that migrations are much faster than the epidemic process, we build up a less dimensional (aggregated) system that allows to study some features of the asymptotic behavior of the original model. In particular, we are able to define global reproduction numbers in the non-spatialized aggregated system that serve to decide the eradication or endemicity of the epidemic in the initial spatially distributed nonautonomous model. Comparing these global reproductive numbers with those corresponding to isolated patches we show that adequate periodic fast migrations can in many cases reverse local endemicity and get global eradication of the epidemic.     

Time-integrated position error accounts for sensorimotor behavior in time-constrained tasks

Several studies have shown that human motor behavior can be successfully described using optimal control theory, which describes behavior by optimizing the trade-off between the subject's effort and performance. This approach predicts that subjects reach the goal exactly at the final time. However, another strategy might be that subjects try to reach the target position well before the final time to avoid the risk of missing the target. To test this, we have investigated whether minimizing the control effort and maximizing the performance is sufficient to describe human motor behavior in time-constrained motor tasks. In addition to the standard model, we postulate a new model which includes an additional cost criterion which penalizes deviations between the position of the effector and the target throughout the trial, forcing arrival on target before the final time. To investigate which model gives the best fit to the data and to see whether that model is generic, we tested both models in two different tasks where subjects used a joystick to steer a ball on a screen to hit a target (first task) or one of two targets (second task) before a final time. Noise of different amplitudes was superimposed on the ball position to investigate the ability of the models to predict motor behavior for different levels of uncertainty. The results show that a cost function representing only a trade-off between effort and accuracy at the end time is insufficient to describe the observed behavior. The new model correctly predicts that subjects steer the ball to the target position well before the final time is reached, which is in agreement with the observed behavior. This result is consistent for all noise amplitudes and for both tasks.     

The relationship between dominance, corticosterone, memory, and food caching in mountain chickadees (Poecile gambeli)

It has been hypothesized that in avian social groups subordinate individuals should maintain more energy reserves than dominants, as an insurance against increased perceived risk of starvation. Subordinates might also have elevated baseline corticosterone levels because corticosterone is known to facilitate fattening in birds. Recent experiments showed that moderately elevated corticosterone levels resulting from unpredictable food supply are correlated with enhanced cache retrieval efficiency and more accurate performance on a spatial memory task. Given the correlation between corticosterone and memory, a further prediction is that subordinates might be more efficient at cache retrieval and show more accurate performance on spatial memory tasks. We tested these predictions in dominant-subordinate pairs of mountain chickadees (Poecile gambeli). Each pair was housed in the same cage but caching behavior was tested individually in an adjacent aviary to avoid the confounding effects of small spaces in which birds could unnaturally and directly influence each other's behavior. In sharp contrast to our hypothesis, we found that subordinate chickadees cached less food, showed less efficient cache retrieval, and performed significantly worse on the spatial memory task than dominants. Although the behavioral differences could have resulted from social stress of subordination, and dominant birds reached significantly higher levels of corticosterone during their response to acute stress compared to subordinates, there were no significant differences between dominants and subordinates in baseline levels or in the pattern of adrenocortical stress response. We find no evidence, therefore, to support the hypothesis that subordinate mountain chickadees maintain elevated baseline corticosterone levels whereas lower caching rates and inferior cache retrieval efficiency might contribute to reduced survival of subordinates commonly found in food-caching parids.