Iain D. Couzin

Effective leadership and decision-making in animal groups on the move

For animals that forage or travel in groups, making movement decisions often depends on social interactions among group members. However, in many cases, few individuals have pertinent information, such as knowledge about the location of a food source, or of a migration route. Using a simple model we show how information can be transferred within groups both without signalling and when group members do not know which individuals, if any, have information. We reveal that the larger the group the smaller the proportion of informed individuals needed to guide the group, and that only a very small proportion of informed individuals is required to achieve great accuracy. We also demonstrate how groups can make consensus decisions, even though informed individuals do not know whether they are in a majority or minority, how the quality of their information compares with that of others, or even whether there are any other informed individuals. Our model provides new insights into the mechanisms of effective leadership and decision-making in biological systems.

From Disorder to Order in Marching Locusts

Recent models from theoretical physics have predicted that mass-migrating animal groups may share group-level properties, irrespective of the type of animals in the group. One key prediction is that as the density of animals in the group increases, a rapid transition occurs from disordered movement of individuals within the group to highly aligned collective motion. Understanding such a transition is crucial to the control of mobile swarming insect pests such as the desert locust. We confirmed the prediction of a rapid transition from disordered to ordered movement and identified a critical density for the onset of coordinated marching in locust nymphs. We also demonstrated a dynamic instability in motion at densities typical of locusts in the field, in which groups can switch direction without external perturbation, potentially facilitating the rapid transfer of directional information.

Self-Organization and Collective Behavior in Vertebrates

Publisher Summary The chapter discusses an emerging area of study: that of applying self-organization theory to mobile vertebrate groups composed of many interacting individuals such as bird flocks, ungulate herds, fish schools, and human crowds in an attempt to improve our understanding of underlying organizational principles. Mathematical modeling is becoming increasingly recognized as an important research tool when studying collective behavior. The chapter presents the interaction dynamics among individuals result in the formation, internal structuring, and collective behaviors of vertebrate groups. The chapter explores the distribution of grouping individuals over larger spatial and temporal scales, and discusses how individual behaviors lead to population-level dynamics. Behavioral differences among individuals within a group may have an important internal structuring influence. By using simulation models, it can be shown how individuals can modify their positions relative to other group members without necessitating information about their current position within the group. In considering self-organization within vertebrate groups it is evident that the organization at one level, for example, that of the group relates to that at higher levels. For example, self-sorting processes that lead to internal structuring within groups also result in population-level patterns when such groups fragment, thus affecting the probability that an individual will be in a group of a given size and composition at any moment in time. These population properties then feed back to the individual interactions by changing the probability of encounters among different members of a population. The chapter concludes that to understand collective behaviors fully, these properties cannot necessarily be considered in isolation.

Collective cognition in animal groups

The remarkable collective action of organisms such as swarming ants, schooling fish and flocking birds has long captivated the attention of artists, naturalists, philosophers and scientists. Despite a long history of scientific investigation, only now are we beginning to decipher the relationship between individuals and group-level properties. This interdisciplinary effort is beginning to reveal the underlying principles of collective decision-making in animal groups, demonstrating how social interactions, individual state, environmental modification and processes of informational amplification and decay can all play a part in tuning adaptive response. It is proposed that important commonalities exist with the understanding of neuronal processes and that much could be learned by considering collective animal behavior in the framework of cognitive science.

Inferring the structure and dynamics of interactions in schooling fish

Determining individual-level interactions that govern highly coordinated motion in animal groups or cellular aggregates has been a long-standing challenge, central to understanding the mechanisms and evolution of collective behavior. Numerous models have been proposed, many of which display realistic-looking dynamics, but nonetheless rely on untested assumptions about how individuals integrate information to guide movement. Here we infer behavioral rules directly from experimental data. We begin by analyzing trajectories of golden shiners (Notemigonus crysoleucas) swimming in two-fish and three-fish shoals to map the mean effective forces as a function of fish positions and velocities. Speeding and turning responses are dynamically modulated and clearly delineated. Speed regulation is a dominant component of how fish interact, and changes in speed are transmitted to those both behind and ahead. Alignment emerges from attraction and repulsion, and fish tend to copy directional changes made by those ahead. We find no evidence for explicit matching of body orientation. By comparing data from two-fish and three-fish shoals, we challenge the standard assumption, ubiquitous in physics-inspired models of collective behavior, that individual motion results from averaging responses to each neighbor considered separately; three-body interactions make a substantial contribution to fish dynamics. However, pairwise interactions qualitatively capture the correct spatial interaction structure in small groups, and this structure persists in larger groups of 10 and 30 fish. The interactions revealed here may help account for the rapid changes in speed and direction that enable real animal groups to stay cohesive and amplify important social information.

Quorum decision-making facilitates information transfer in fish shoals

Despite the growing interest in collective phenomena such as “swarm intelligence” and “wisdom of the crowds,” little is known about the mechanisms underlying decision-making in vertebrate animal groups. How do animals use the behavior of others to make more accurate decisions, especially when it is not possible to identify which individuals possess pertinent information? One plausible answer is that individuals respond only when they see a threshold number of individuals perform a particular behavior. Here, we investigate the role of such “quorum responses” in the movement decisions of fish (three-spine stickleback, Gasterosteus aculeatus). We show that a quorum response to conspecifics can explain how sticklebacks make collective movement decisions, both in the absence and presence of a potential predation risk. Importantly our experimental work shows that a quorum response can reduce the likelihood of amplification of nonadaptive following behavior. Whereas the traveling direction of solitary fish was strongly influenced by a single replica conspecific, the replica was largely ignored by larger groups of four or eight sticklebacks under risk, and the addition of a second replica was required to exert influence on the movement decisions of such groups. Model simulations further predict that quorum responses by fish improve the accuracy and speed of their decision-making over that of independent decision-makers or those using a weak linear response. This study shows that effective and accurate information transfer in groups may be gained only through nonlinear responses of group members to each other, thus highlighting the importance of quorum decision-making.

Context-dependent group size choice in fish

The costs and benefits of group membership vary with the size of groups, and individuals are expected to modify their choice of groups in response to ecological factors such as food availability and predation risk. We experimentally examined context-dependent group size choice in a shoaling fish, the banded killifish, Fundulus diaphanus, by using nondirectional odour cues to simulate a food source or a successful attack by a predator (food or alarm treatments) in the laboratory. Group sizes were significantly smaller in the food treatment and larger in the alarm treatment than in control trials. When presented with food and alarm cues together, fish formed groups that were larger than control groups but smaller than those seen with alarm cues alone. These results are consistent with theoretical predictions based on the known benefits and costs of grouping and with previous laboratory work examining the individual shoal choice behaviour of single fish. To examine possible mechanisms of group formation, we developed an individual-based model of shoaling behaviour in which simulated fish were allowed to modify the area over which they interacted with neighbouring individuals. Group size distributions produced by the model were a good approximation of our experimental data. We suggest that local behavioural interaction rules of this type are a potential mechanism by which fish may individually adjust grouping behaviour without requiring extensive information on the position and movement of all possible shoalmates.

Self-organized lane formation and optimized traffic flow in army ants

We show how the movement rules of individual ants on trails can lead to a collective choice of direction and the formation of distinct traffic lanes that minimize congestion. We develop and evaluate the results of a new model with a quantitative study of the behaviour of the army ant Eciton burchelli. Colonies of this species have up to 200 000 foragers and transport more than 3000 prey items per hour over raiding columns that exceed 100 m. It is an ideal species in which to test the predictions of our model because it forms pheromone trails that are densely populated with very swift ants. The model explores the influences of turning rates and local perception on traffic flow. The behaviour of real army ants is such that they occupy the specific region of parameter space in which lanes form and traffic flow is maximized.

Leadership, consensus decision making and collective behaviour in humans

This paper reviews the literature on leadership in vertebrate groups, including recent work on human groups, before presenting the results of three new experiments looking at leadership and decision making in small and large human groups. In experiment 1, we find that both group size and the presence of uninformed individuals can affect the speed with which small human groups (eight people) decide between two opposing directional preferences and the likelihood of the group splitting. In experiment 2, we show that the spatial positioning of informed individuals within small human groups (10 people) can affect the speed and accuracy of group motion. We find that having a mixture of leaders positioned in the centre and on the edge of a group increases the speed and accuracy with which the group reaches their target. In experiment 3, we use large human crowds (100 and 200 people) to demonstrate that the trends observed from earlier work using small human groups can be applied to larger crowds. We find that only a small minority of informed individuals is needed to guide a large uninformed group. These studies build upon important theoretical and empirical work on leadership and decision making in animal groups.

Uninformed Individuals Promote Democratic Consensus in Animal Groups

Uninformed individuals inhibit extremism and enforce fair representation during collective decision-making. Conflicting interests among group members are common when making collective decisions, yet failure to achieve consensus can be costly. Under these circumstances individuals may be susceptible to manipulation by a strongly opinionated, or extremist, minority. It has previously been argued, for humans and animals, that social groups containing individuals who are uninformed, or exhibit weak preferences, are particularly vulnerable to such manipulative agents. Here, we use theory and experiment to demonstrate that, for a wide range of conditions, a strongly opinionated minority can dictate group choice, but the presence of uninformed individuals spontaneously inhibits this process, returning control to the numerical majority. Our results emphasize the role of uninformed individuals in achieving democratic consensus amid internal group conflict and informational constraints.