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Swarming, schooling, milling: phase diagram of a data-driven fish school model

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Swarming, schooling, milling: phase diagram of a data-driven fish school model
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We determine the basic phase diagram of the fish school model derived from data by Gautrais et al (2012 PLoS Comput. Biol. 8 e1002678), exploring its parameter space beyond the parameter values determined experimentally on groups of barred flagtails (Kuhlia mugil) swimming in a shallow tank. A modified model is studied alongside the original one, in which an additional frontal preference is introduced in the stimulus/response function to account for the angular weighting of interactions. Our study, mostly limited to groups of moderate size (in the order of 100 individuals), focused not only on the transition to schooling induced by increasing the swimming speed, but also on the conditions under which a school can exhibit milling dynamics and the corresponding behavioural transitions. We show the existence of a transition region between milling and schooling, in which the school exhibits multistability and intermittence between schooling and milling for the same combination of individual parameters. We also show that milling does not occur for arbitrarily large groups, mainly due to a distance dependence interaction of the model and information propagation delays in the school, which cause conflicting reactions for large groups. We finally discuss the biological significance of our findings, especially the dependence of behavioural transitions on social interactions, which were reported by Gautrais et al to be adaptive in the experimental conditions.
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Transcript: English(auto-generated)
Swarming, schooling, milling, phase diagram of a data-driven fish school model. Fish schools are able to display some amazing collective patterns in nature. These patterns arise naturally from their interactions and are linked to behaviors such as migration, feeding, predator vision, etc.
In this article, we use a previously developed model to understand the amount of naturally and especially after proposing a different angular preference in the directions.
This model was developed for this species called the musu and consists in a persistent turning walker with a fixed swimming speed. This means that fish interactions and noise affect the angular velocity of the fish. The interactions are calculated for every pair in the first shell of the Voronoi tessellation and consist in attraction and alignment turn.
Experimental data review the forward angular preference which led us to investigate the effects of this new yet simple modulation of interactional form. The first result obtained was given by the polarization of the school for different
velocities where we can see that in both cases we have a transition from a swarming to a schooling state as the speed of the school increases. Doing a parameter scan of the alignment and attraction parameters in non-dimensionalized equations while analyzing the polarization and the absolute values of the normalized
angular momentum, we could detect that both choices display a transition from swarming to schooling. We can also see that with the angular preference the schooling behavior has a dependence on the attraction parameter due to the fact that the system now exhibits a milling behavior
which was not found without the angular preference. Analyzing the average neighbor distance between fish we expected these values to diverge when the attraction parameter tends to zero and to decay when increased. However, the parameter space with the angular preference displayed a second reach as the
attraction parameter was increased. We can define different colors for each of these three behaviors, red for schooling, green for high average distance, and blue for milling to plot this behavioral map. In region 1 we can see the following schooling behavior.
Simulations on region 2 show a very good example of a fish mill. We can also see the transition between schooling and milling region where we can see spontaneous changes between both states. In the high average distance region we can see that fish organize themselves into files.
This is a consequence of the frontal preference and the fact that attraction dominates interactions while allowing anti-parallel configurations. We have also studied the transition between the schooling and milling state and verified
it follows a simple functional form, which is independent of the fish speed. A group size impact study was also done for a fixed attraction parameter and for different alignment values. We can see that there is a minimum number of 60 fish to get a milling behavior. And also that the model is not able to produce arbitrarily large fish vortexes.
In conclusion, we have shown that the relative weights of the attraction alignment interactions play a key role in the emerging collective states that emerge in the school level. Depending on the magnitude of the attraction and alignment of fish to their neighbors, different collective states can be reached by the school.