Jolle Wolter Jolles

Dominance, pair bonds and boldness determine social-foraging tactics in rooks, Corvus frugilegus

Socially foraging animals can search for resources themselves (produce) or exploit the discoveries made by others (scrounge). The extensive literature on producerescrounger dynamics has mainly focused on scramble competition over readily accessible resources, thereby largely neglecting the variety of scrounging techniques individuals may use as well as the role of investment in food handling. Furthermore, although individual differences in boldness and social factors such as dominance have been described to influence foraging tactics, their potential interplay and effect in foraging contexts beyond the conventional producerescrounger game remains unclear. We investigated the relationship between social-foraging tactic use and dominance, pair bonds and boldness in a foraging experiment focused on food handling and alternative scrounging tactics. We conducted a producerescrounger experiment in a captive group of rooks in which individuals could produce by pulling up baited strings, or scrounge by retrieving fallen food items or joining a producer. There were three key findings: (1) dominant rooks adopted the producer tactic more often and more successfully than subordinates; (2) producing and scrounging by tolerance led to mixed benefits to paired birds; (3) bold birds scrounged by retrieving more often than shy birds. Importantly, individuals were highly consistent in their tactic use across conditions differing in food availability. Our study highlights the importance of taking both social factors and boldness (heterogeneity) into account when studying social-foraging dynamics and offers empirical data on food handling and alternative scrounging tactics that can be used to extend current models and experiments on social foraging.

Male and female Wistar rats differ in decision-making performance in a rodent version of the Iowa Gambling Task

The Iowa Gambling Task (IGT) measures decision-making processes by simulating real-life decisions involving reward, punishment, and uncertainty of outcomes. In humans, men show more choices for the advantageous option than women. Here, we investigated sex differences in a rat model of the IGT (r-IGT). In our r-IGT mildly food-deprived rats learn to differentiate a long-term advantageous arm from a long-term disadvantageous arm differing in frequency and amount of sugar pellets as well as unpalatable but not uneatable quinine-treated sugar pellets. We also used a T-maze discrimination procedure in which rats learn to differentiate a high from a low reward arm to further explore sex differences in reward-related decision-making. In line with human data, male rats showed a stronger task progression of choices for the advantageous option than female rats. Furthermore, male rats showed more win-stay and less lose-shift behaviour in the advantageous arm as the task progressed than female rats. Whilst both male and female rats had a stronger preference for the high over the small reward arm in the T-maze, males increased this preference over sessions, whilst females did not. These data are discussed in relation to sex differences in processing rewards and punishments.

The role of social attraction and its link with boldness in the collective movements of three-spined sticklebacks

Social animals must time and coordinate their behaviour to ensure the benefits of grouping, resulting in collective movements and the potential emergence of leaders and followers. However, individuals often differ consistently from one another in how they cope with their environment, a phenomenon known as animal personality, which may affect how individuals use coordination rules and requiring them to compromise. Here we tracked the movements of pairs of three-spined sticklebacks, Gasterosteus aculeatus, separated by a transparent partition that allowed them to observe and interact with one another in a context containing cover. Individuals differed consistently in their tendency to approach their partners compartment during collective movements. The strength of this social attraction was positively correlated with the behavioural coordination between members of a pair but was negatively correlated with an individuals tendency to lead. Social attraction may form part of a broader behavioural syndrome as it was predicted by the boldness of an individual, measured in isolation prior to the observation of pairs, and by the boldness of the partner. We found that bolder fish, and those paired with bolder partners, tended to approach their partners compartment less closely. These findings provide important insights into the mechanisms that govern the dynamics and functioning of social groups and the emergence and maintenance of consistent behavioural differences.

Social modulation of decision-making: a cross-species review

Taking decisions plays a pivotal role in daily life and comprises a complex process of assessing and weighing short-term and long-term costs and benefits of competing actions. Decision-making has been shown to be affected by factors such as sex, age, genotype, and personality. Importantly, also the social environment affects decisions, both via social interactions (e.g., social learning, cooperation and competition) and social stress effects. Although everyone is aware of this social modulating role on daily life decisions, this has thus far only scarcely been investigated in human and animal studies. Furthermore, neuroscientific studies rarely discuss social influence on decision-making from a functional perspective such as done in behavioral ecology studies. Therefore, the first aim of this article is to review the available data of the influence of the social context on decision-making both from a causal and functional perspective, drawing on animal and human studies. Also, there is currently still a gap between decision-making in real life where influences of the social environment are extensive, and decision-making as measured in the laboratory, which is often done without any (deliberate) social influences. However, methods are being developed to bridge this gap. Therefore, the second aim of this review is to discuss these methods and ways in which this gap can be increasingly narrowed. We end this review by formulating future research questions.

Consistent Individual Differences Drive Collective Behavior and Group Functioning of Schooling Fish

Summary The ubiquity of consistent inter-individual differences in behavior (“animal personalities”) [1, 2] suggests that they might play a fundamental role in driving the movements and functioning of animal groups [3, 4], including their collective decision-making, foraging performance, and predator avoidance. Despite increasing evidence that highlights their importance [5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16], we still lack a unified mechanistic framework to explain and to predict how consistent inter-individual differences may drive collective behavior. Here we investigate how the structure, leadership, movement dynamics, and foraging performance of groups can emerge from inter-individual differences by high-resolution tracking of known behavioral types in free-swimming stickleback (Gasterosteus aculeatus) shoals. We show that individual’s propensity to stay near others, measured by a classic “sociability” assay, was negatively linked to swim speed across a range of contexts, and predicted spatial positioning and leadership within groups as well as differences in structure and movement dynamics between groups. In turn, this trait, together with individual’s exploratory tendency, measured by a classic “boldness” assay, explained individual and group foraging performance. These effects of consistent individual differences on group-level states emerged naturally from a generic model of self-organizing groups composed of individuals differing in speed and goal-orientedness. Our study provides experimental and theoretical evidence for a simple mechanism to explain the emergence of collective behavior from consistent individual differences, including variation in the structure, leadership, movement dynamics, and functional capabilities of groups, across social and ecological scales. In addition, we demonstrate individual performance is conditional on group composition, indicating how social selection may drive behavioral differentiation between individuals.

Heterogeneous structure in mixed-species corvid flocks in flight

Flocks of birds in flight represent a striking example of collective behaviour. Models of self-organization suggest that repeated interactions among individuals following simple rules can generate the complex patterns and coordinated movements exhibited by flocks. However, such models often assume that individuals are identical and interchangeable, and fail to account for individual differences and social relationships among group members. Here, we show that heterogeneity resulting from species differences and social structure can affect flock spatial dynamics. Using high-resolution photographs of mixed flocks of jackdaws, Corvus monedula, and rooks, Corvus frugilegus, we show that birds preferentially associated with conspecifics and that, like high-ranking members of single-species groups, the larger and more socially dominant rooks positioned themselves near the leading edge of flocks. Neighbouring birds showed closer directional alignment if they were of the same species, and neighbouring jackdaws in particular flew very close to one another. Moreover, birds of both species often flew especially close to a single same-species neighbour, probably reflecting the monogamous pair bonds that characterize these corvid social systems. Together, our findings demonstrate that the characteristics of individuals and their social systems are likely to result in preferential associations that critically influence flock structure. 2013 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved. How do large aggregations of individuals, each of which may differ in its preferred outcome, coordinate their movements? The spectacular displays of flocking birds led the naturalist Edmund Selous (1931) to postulate a role for ‘thought transference’, but recent advances have begun to unravel the mysteries of collective movement without appealing to the supernatural (Couzin & Krause 2003; Conradt & Roper 2005; Sumpter 2006). Models of selforganizing systems suggest that repeated interactions among individuals following simple rules can generate complex patterns and coordinated group movements. Models of agents following simple rules of (1) long-range attraction to group members, (2) shortrange repulsion and (3) alignment between close neighbours have generated realistic representations of collective animal movements (reviewed in Sumpter 2006; Petit & Bon 2010). However, empirical verification of their assumptions remains scarce and largelyconfined to model systems such as starlings, Sturnus vulgaris

Recent social conditions affect boldness repeatability in individual sticklebacks

Animal personalities are ubiquitous across the animal kingdom and have been shown both to influence individual behaviour in the social context and to be affected by it. However, little attention has been paid to possible carryover effects of social conditions on personality expression, especially when individuals are alone. Here we investigated how the recent social context affected the boldness and repeatability of three-spined sticklebacks, Gasterosteus aculeatus, during individual assays. We housed fish either solitarily, solitarily part of the time or socially in groups of four, and subjected them twice to a risk-taking task. The social conditions had a large effect on boldness repeatability, with fish housed solitarily before the trials showing much higher behavioural repeatability than fish housed socially, for which repeatability was not significant. Social conditions also had a temporal effect on the boldness of the fish, with only fish housed solitarily taking more risks during the first than the second trial. These results show that recent social conditions can thus affect the short-term repeatability of behaviour and obfuscate the expression of personality even in later contexts when individuals are alone. This finding highlights the need to consider social housing conditions when designing personality studies and emphasizes the important link between animal personality and the social context by showing the potential role of social carryover effects.

Male Wistar rats show individual differences in an animal model of conformity.

Conformity refers to the act of changing one’s behaviour to match that of others. Recent studies in humans have shown that individual differences exist in conformity and that these differences are related to differences in neuronal activity. To understand the neuronal mechanisms in more detail, animal tests to assess conformity are needed. Here, we used a test of conformity in rats that has previously been evaluated in female, but not male, rats and assessed the nature of individual differences in conformity. Male Wistar rats were given the opportunity to learn that two diets differed in palatability. They were subsequently exposed to a demonstrator that had consumed the less palatable food. Thereafter, they were exposed to the same diets again. Just like female rats, male rats decreased their preference for the more palatable food after interaction with demonstrator rats that had eaten the less palatable food. Individual differences existed for this shift, which were only weakly related to an interaction between their own initial preference and the amount consumed by the demonstrator rat. The data show that this conformity test in rats is a promising tool to study the neurobiology of conformity.

Food intake rates of inactive fish are positively linked to boldness in three-spined sticklebacks Gasterosteus aculeatus

To investigate the link between personality and maximum food intake of inactive individuals, food‐deprived three‐spined sticklebacks Gasterosteus aculeatus at rest in their home compartments were provided with ad libitum prey items. Bolder individuals ate considerably more than shyer individuals, even after accounting for body size, while sociability did not have an effect. These findings support pace‐of‐life theory predicting that life‐history strategies are linked to boldness.

Seasonal changes in neophobia and its consistency in rooks: the effect of novelty type and dominance position.

Neophobia, or the fear of novelty, may offer benefits to animals by limiting their exposure to unknown danger, but can also impose costs by preventing the exploration of potential resources. The costs and benefits of neophobia may vary throughout the year if predation pressure, resource distribution or conspecific competition changes seasonally. Despite such variation, neophobia levels are often assumed to be temporally and individually stable. Whether or not neophobia expression changes seasonally and fluctuates equally for all individuals is crucial to understanding the drivers, consequences and plasticity of novelty avoidance. We investigated seasonal differences and individual consistency in the motivation and novelty responses of a captive group of rooks, Corvus frugilegus, a seasonally breeding, colonial species of corvid that is known for being neophobic. We tested the group around novel objects and novel people to determine whether responses generalized across novelty types, and considered whether differences in dominance could influence the social risk of approaching unknown stimuli. We found that the groups level of object neophobia was stable year-round, but individuals were not consistent between seasons, despite being consistent within seasons. In contrast, the groups avoidance of novel people decreased during the breeding season, and individuals were consistent year-round. Additionally, although subordinate birds were more likely to challenge dominants during the breeding season, this social risk taking did not translate to greater novelty approach. Since seasonal variation and individual consistency varied differently towards each novelty type, responses towards novel objects and people seem to be governed by different mechanisms. Such a degree of fluctuation has consequences for other individually consistent behaviours often measured within the nonhuman personality literature.