In many species, group cohesion may be the result of a compromise between opposing forces (e.g. predator avoidance and competition for food). However, little empirical data exists on the dynamics of group cohesion. We present moment-to-moment positional data on zebrafish shoals and analyze temporal changes in inter-individual distances. We demonstrate that the distance between shoal members does not settle at any given value, as has previously been assumed, but oscillates with a period between 5 and 15 s.
KeywordsShoaling; Schooling; Zebrafish; Oscillations Many species, from ants to humans, live in groups and interact with each other [13,14]. Members of groups often face opposing forces: some factors may favor tighter groups and others favor more dispersed groups. For example, numerous group-forming species must balance the requirements of efficient foraging with protection from predation: foraging efficiency increases with decreasing group density [10,12] but protection from predation decreases [4,6]. Many researchers have argued that as a result of such opposing forces, group density should settle at a species-specific optimal value. Whilst several models have been developed to take such opposing forces into account and to explain how an optimal group density or size are achieved (e.g. Refs. [16,7]), the temporal dynamics of group density changes, if they occur at all, have been ignored. Most previous experimental studies of group behavior have averaged the distances between group members over several observation periods, or considered only the nearest neighbors of each individual (e.g. Ref. [8]). Thus, the data to validate or disprove the existence of an optimal group density or its temporal stability do not exist.Here, using custom software [5], we examined high temporal resolution data on the positions of free-swimming zebrafish (Daniorerio). We focused on the distances between individuals within a group (AvD) and examined how they change over time. Our initial examination of the data suggested that there may be regular oscillations in shoal density, as measured by the average distance between fish, and we investigated whether these oscillations were robust and whether they had a characteristic frequency. We show that zebrafish do not settle on an average distance from conspecifics in the shoal, but that shoal density fluctuates with a characteristic frequency. To examine shoaling preferences, 3 groups of 16 zebrafish each were placed in a 91-cm diameter circular tank and filmed from above for 30min per session for 6 sessions. Using custom software described in detail elsewhere [5], we extracted from the videos of the experiments the average distance in cm between any two fish (AvD). AvD is the average of the inter-individual distances (IIDs), which are the average distance of a given fish from all the other fish in the enclosure.Five 1-min long sections of the sessions (at 5, 10, 15, 20, and 25 min from the start of the session) were coded at 1 s intervals (one frame per second). Thus, each datase...