Collective motion phenomena in large groups of social organisms have long fascinated the observer, especially in cases, such as bird flocks or fish schools, where large-scale highly coordinated actions emerge in the absence of obvious leaders. However, the mechanisms involved in this self-organized behavior are still poorly understood, because the individual-level interactions underlying them remain elusive. Here, we demonstrate the power of a bottom-up methodology to build models for animal group motion from data gathered at the individual scale. Using video tracks of fish shoal in a tank, we show how a careful, incremental analysis at the local scale allows for the determination of the stimulus/response function governing an individual's moving decisions. We find in particular that both positional and orientational effects are present, act upon the fish turning speed, and depend on the swimming speed, yielding a novel schooling model whose parameters are all estimated from data. Our approach also leads to identify a density-dependent effect that results in a behavioral change for the largest groups considered. This suggests that, in confined environment, the behavioral state of fish and their reaction patterns change with group size. We debate the applicability, beyond the particular case studied here, of this novel framework for deciphering interactions in moving animal groups.
Swarms of insects, schools of fish and flocks of birds display an impressive variety of collective patterns that emerge from local interactions among group members. These puzzling phenomena raise a variety of questions about the behavioral rules that govern the coordination of individuals' motions and the emergence of large-scale patterns. While numerous models have been proposed, there is still a strong need for detailed experimental studies to foster the biological understanding of such collective motion. Here, we use data recorded on fish barred flagtails moving in groups of increasing sizes in a water tank to demonstrate the power of an incremental methodology for building a fish behavior model completely based on interactions with the physical environment and neighboring fish. In contrast to previous works, our model revealed an implicit balancing of neighbors position and orientation on the turning speed of fish, an unexpected transition between shoaling and schooling induced by a change in the swimming speed, and a group-size effect which results in a decrease of social interactions among fish as density increases. An important feature of this model lies in its ability to allow a large palette of adaptive patterns with a great economy of means.