Unravelling the origins of boldness behaviour: a common garden experiment with cavefish (Barbatula barbatula).

Many animals show an aversion to bright, open spaces, with significant variability seen across species, populations and individuals within populations. Although there is much interest in the underlying causes of this behaviour, few studies have been able to systematically isolate the role of heritable and environmental effects. Here, we addressed this gap using a common garden experiment with cavefish. Specifically, we bred and cross-bred cave loaches (Barbatula barbatula), Europe's only known cavefish, in the laboratory, raised the offspring in complete darkness or normal light conditions, and studied their light avoidance behaviour. Cavefish spent much more time in a light area and ventured further out, while surface fish spent considerable time in risk-assessment behaviour between the light and dark areas. Hybrids behaved most similarly to cavefish. Light treatment and eye quality and lens size only had a modest effect. Our results suggest light avoidance behaviour of cavefish has a heritable basis and is fundamentally linked to increased boldness rather than reduced vision, which is likely adaptive given the complete lack of macropredators in the cave environment. Our study provides novel experimental insights into the behavioural divergence of cavefish and contributes to our broader understanding of the evolution of boldness and behavioural adaptation.

The emergence and development of behavioral individuality in clonal fish.

Behavioral individuality is a ubiquitous phenomenon in animal populations, yet the origins and developmental trajectories of individuality, especially very early in life, are still a black box. Using a high-resolution tracking system, we mapped the behavioral trajectories of genetically identical fish (Poecilia formosa), separated immediately after birth into identical environments, over the first 10 weeks of their life at 3 s resolution. We find that (i) strong behavioral individuality is present at the very first day after birth, (ii) behavioral differences at day 1 of life predict behavior up to at least 10 weeks later, and (iii) patterns of individuality strengthen gradually over developmental time. Our results establish a null model for how behavioral individuality can develop in the absence of genetic and environmental variation and provide experimental evidence that later-in-life individuality can be strongly shaped by factors pre-dating birth like maternal provisioning, epigenetics and pre-birth developmental stochasticity.

Vocally mediated consensus decisions govern mass departures from jackdaw roosts.

In the early morning, large groups of up to hundreds or even thousands of roosting birds, sometimes comprising the entire roost population, often take off together in sudden mass departures. These departures commonly occur in low-light conditions and structurally complex habitats where access to visual cues is likely to be restricted. Roosting birds are often highly vocal, leading us to hypothesise that vocalisations, which can propagate over large distances, could provide a means of enabling individuals to agree on when to depart - that is to establish a consensus1 - and thus coordinate the timing of mass movements. Investigations of the role of acoustic signals in coordinating collective decisions have been limited to honeybees2 and relatively small vertebrate groups (<50 individuals)3-5 and have rarely included experimental validation2,3. Here, by combining field recordings with a large-scale experimental manipulation, we show that jackdaws (Corvus monedula) use vocalisations to coordinate mass departures from winter roosts. This provides empirical evidence for vocally-mediated consensus decision-making in large vertebrate groups.

Open Hardware in Science: The Benefits of Open Electronics.

Openly shared low-cost electronic hardware applications, known as open electronics, have sparked a new open-source movement, with much untapped potential to advance scientific research. Initially designed to appeal to electronic hobbyists, open electronics have formed a global "maker" community and are increasingly used in science and industry. In this perspective article, we review the current costs and benefits of open electronics for use in scientific research ranging from the experimental to the theoretical sciences. We discuss how user-made electronic applications can help (I) individual researchers, by increasing the customization, efficiency, and scalability of experiments, while improving data quantity and quality; (II) scientific institutions, by improving access to customizable high-end technologies, sustainability, visibility, and interdisciplinary collaboration potential; and (III) the scientific community, by improving transparency and reproducibility, helping decouple research capacity from funding, increasing innovation, and improving collaboration potential among researchers and the public. We further discuss how current barriers like poor awareness, knowledge access, and time investments can be resolved by increased documentation and collaboration, and provide guidelines for academics to enter this emerging field. We highlight that open electronics are a promising and powerful tool to help scientific research to become more innovative and reproducible and offer a key practical solution to improve democratic access to science.

The Potential for Physiological Performance Curves to Shape Environmental Effects on Social Behavior.

As individual animals are exposed to varying environmental conditions, phenotypic plasticity will occur in a vast array of physiological traits. For example, shifts in factors such as temperature and oxygen availability can affect the energy demand, cardiovascular system, and neuromuscular function of animals that in turn impact individual behavior. Here, we argue that nonlinear changes in the physiological traits and performance of animals across environmental gradients-known as physiological performance curves-may have wide-ranging effects on the behavior of individual social group members and the functioning of animal social groups as a whole. Previous work has demonstrated how variation between individuals can have profound implications for socially living animals, as well as how environmental conditions affect social behavior. However, the importance of variation between individuals in how they respond to changing environmental conditions has so far been largely overlooked in the context of animal social behavior. First, we consider the broad effects that individual variation in performance curves may have on the behavior of socially living animals, including: (1) changes in the rank order of performance capacity among group mates across environments; (2) environment-dependent changes in the amount of among- and within-individual variation, and (3) differences among group members in terms of the environmental optima, the critical environmental limits, and the peak capacity and breadth of performance. We then consider the ecological implications of these effects for a range of socially mediated phenomena, including within-group conflict, within- and among group assortment, collective movement, social foraging, predator-prey interactions and disease and parasite transfer. We end by outlining the type of empirical work required to test the implications for physiological performance curves in social behavior.

Fisheries-induced changes of shoaling behaviour: mechanisms and potential consequences.

We outline key mechanisms by which fishing can change the shoaling tendency and collective behaviour of exploited species - an issue that is rarely considered and poorly understood. We highlight potential consequences for fish populations and food webs, and discuss possible repercussions for fisheries and conservation strategies.

Group-level patterns emerge from individual speed as revealed by an extremely social robotic fish.

Understanding the emergence of collective behaviour has long been a key research focus in the natural sciences. Besides the fundamental role of social interaction rules, a combination of theoretical and empirical work indicates individual speed may be a key process that drives the collective behaviour of animal groups. Socially induced changes in speed by interacting animals make it difficult to isolate the effects of individual speed on group-level behaviours. Here, we tackled this issue by pairing guppies with a biomimetic robot. We used a closed-loop tracking and feedback system to let a robotic fish naturally interact with a live partner in real time, and programmed it to strongly copy and follow its partner's movements while lacking any preferred movement speed or directionality of its own. We show that individual differences in guppies' movement speed were highly repeatable and in turn shaped key collective patterns: a higher individual speed resulted in stronger leadership, lower cohesion, higher alignment and better temporal coordination of the pairs. By combining the strengths of individual-based models and observational work with state-of-the-art robotics, we provide novel evidence that individual speed is a key, fundamental process in the emergence of collective behaviour.

Schistocephalus parasite infection alters sticklebacks' movement ability and thereby shapes social interactions.

Parasitism is ubiquitous in the animal kingdom. Although many fundamental aspects of host-parasite relationships have been unravelled, few studies have systematically investigated how parasites affect organismal movement. Here we combine behavioural experiments of Schistocephalus solidus infected sticklebacks with individual-based simulations to understand how parasitism affects individual movement ability and thereby shapes social interaction patterns. High-resolution tracking revealed that infected fish swam, accelerated, and turned more slowly than did non-infected fish, and tended to be more predictable in their movements. Importantly, the strength of these effects increased with increasing parasite load (proportion of body weight), with more heavily infected fish showing larger changes and impairments in behaviour. When grouped, pairs of infected fish moved more slowly, were less cohesive, less aligned, and less temporally coordinated than non-infected pairs, and mixed pairs were primarily led by the non-infected fish. These social patterns also emerged in simulations of self-organised groups composed of individuals differing similarly in speed and turning tendency, suggesting infection-induced changes in mobility and manoeuvrability may drive collective outcomes. Together, our results demonstrate how infection with a complex life-cycle parasite affects the movement ability of individuals and how this in turn shapes social interaction patterns, providing important mechanistic insights into the effects of parasites on host movement dynamics.

The Role of Individual Heterogeneity in Collective Animal Behaviour.

Social grouping is omnipresent in the animal kingdom. Considerable research has focused on understanding how animal groups form and function, including how collective behaviour emerges via self-organising mechanisms and how phenotypic variation drives the behaviour and functioning of animal groups. However, we still lack a mechanistic understanding of the role of phenotypic variation in collective animal behaviour. Here we present a common framework to quantify individual heterogeneity and synthesise the literature to systematically explain and predict its role in collective behaviour across species, contexts, and traits. We show that individual heterogeneity provides a key intermediary mechanism with broad consequences for sociality (e.g., group structure, functioning), ecology (e.g., response to environmental change), and evolution. We also outline a roadmap for future research.

Repeatable group differences in the collective behaviour of stickleback shoals across ecological contexts.

Establishing how collective behaviour emerges is central to our understanding of animal societies. Previous research has highlighted how universal interaction rules shape collective behaviour, and that individual differences can drive group functioning. Groups themselves may also differ considerably in their collective behaviour, but little is known about the consistency of such group variation, especially across different ecological contexts that may alter individuals' behavioural responses. Here, we test if randomly composed groups of sticklebacks differ consistently from one another in both their structure and movement dynamics across an open environment, an environment with food, and an environment with food and shelter. Based on high-resolution tracking data of the free-swimming shoals, we found large context-associated changes in the average behaviour of the groups. But despite these changes and limited social familiarity among group members, substantial and predictable behavioural differences between the groups persisted both within and across the different contexts (group-level repeatability): some groups moved consistently faster, more cohesively, showed stronger alignment and/or clearer leadership than other groups. These results suggest that among-group heterogeneity could be a widespread feature in animal societies. Future work that considers group-level variation in collective behaviour may help understand the selective pressures that shape how animal collectives form and function.

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

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 [5-16], 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.

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 group's level of object neophobia was stable year-round, but individuals were not consistent between seasons, despite being consistent within seasons. In contrast, the group's 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.

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 partner's 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 individual's 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 partner's 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.

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.