Edmondo Silvestri

Statistical mechanics for natural flocks of birds

Flocking is a typical example of emergent collective behavior, where interactions between individuals produce collective patterns on the large scale. Here we show how a quantitative microscopic theory for directional ordering in a flock can be derived directly from field data. We construct the minimally structured (maximum entropy) model consistent with experimental correlations in large flocks of starlings. The maximum entropy model shows that local, pairwise interactions between birds are sufficient to correctly predict the propagation of order throughout entire flocks of starlings, with no free parameters. We also find that the number of interacting neighbors is independent of flock density, confirming that interactions are ruled by topological rather than metric distance. Finally, by comparing flocks of different sizes, the model correctly accounts for the observed scale invariance of long-range correlations among the fluctuations in flight direction.

Social interactions dominate speed control in poising natural flocks near criticality.

Significance The coherent flight of bird flocks is one of nature’s most impressive aerial displays. Beyond the fact that thousands of birds fly, on average, with the same velocity, quantitative observations show that small deviations of individual birds from this average are correlated across the entire flock. By learning minimally structured models from field data, we show that these long-ranged correlations are consistent with local interactions among neighboring birds, but only because the parameters of the flock are tuned to special values, mathematically equivalent to a critical point in statistical mechanics. Being in this critical regime allows information to propagate almost without loss throughout the flock, while keeping the variance of individual velocities small. Flocks of birds exhibit a remarkable degree of coordination and collective response. It is not just that thousands of individuals fly, on average, in the same direction and at the same speed, but that even the fluctuations around the mean velocity are correlated over long distances. Quantitative measurements on flocks of starlings, in particular, show that these fluctuations are scale-free, with effective correlation lengths proportional to the linear size of the flock. Here we construct models for the joint distribution of velocities in the flock that reproduce the observed local correlations between individuals and their neighbors, as well as the variance of flight speeds across individuals, but otherwise have as little structure as possible. These minimally structured or maximum entropy models provide quantitative, parameter-free predictions for the spread of correlations throughout the flock, and these are in excellent agreement with the data. These models are mathematically equivalent to statistical physics models for ordering in magnets, and the correct prediction of scale-free correlations arises because the parameters—completely determined by the data—are in the critical regime. In biological terms, criticality allows the flock to achieve maximal correlation across long distances with limited speed fluctuations.

Collective Behaviour without Collective Order in Wild Swarms of Midges

Collective behaviour is a widespread phenomenon in biology, cutting through a huge span of scales, from cell colonies up to bird flocks and fish schools. The most prominent trait of collective behaviour is the emergence of global order: individuals synchronize their states, giving the stunning impression that the group behaves as one. In many biological systems, though, it is unclear whether global order is present. A paradigmatic case is that of insect swarms, whose erratic movements seem to suggest that group formation is a mere epiphenomenon of the independent interaction of each individual with an external landmark. In these cases, whether or not the group behaves truly collectively is debated. Here, we experimentally study swarms of midges in the field and measure how much the change of direction of one midge affects that of other individuals. We discover that, despite the lack of collective order, swarms display very strong correlations, totally incompatible with models of non-interacting particles. We find that correlation increases sharply with the swarms density, indicating that the interaction between midges is based on a metric perception mechanism. By means of numerical simulations we demonstrate that such growing correlation is typical of a system close to an ordering transition. Our findings suggest that correlation, rather than order, is the true hallmark of collective behaviour in biological systems.

Spatially balanced topological interaction grants optimal cohesion in flocking models

Models of self-propelled particles (SPPs) are an indispensable tool to investigate collective animal behaviour. Originally, SPP models were proposed with metric interactions, where each individual coordinates with neighbours within a fixed metric radius. However, recent experiments on bird flocks indicate that interactions are topological: each individual interacts with a fixed number of neighbours, irrespective of their distance. It has been argued that topological interactions are more robust than metric ones against external perturbations, a significant evolutionary advantage for systems under constant predatory pressure. Here, we test this hypothesis by comparing the stability of metric versus topological SPP models in three dimensions. We show that topological models are more stable than metric ones. We also show that a significantly better stability is achieved when neighbours are selected according to a spatially balanced topological rule, namely when interacting neighbours are evenly distributed in angle around the focal individual. Finally, we find that the minimal number of interacting neighbours needed to achieve fully stable cohesion in a spatially balanced model is compatible with the value observed in field experiments on starling flocks.

Flocking and Turning: a New Model for Self-organized Collective Motion

Birds in a flock move in a correlated way, resulting in large polarization of velocities. A good understanding of this collective behavior exists for linear motion of the flock. Yet observing actual birds, the center of mass of the group often turns giving rise to more complicated dynamics, still keeping strong polarization of the flock. Here we propose novel dynamical equations for the collective motion of polarized animal groups that account for correlated turning including solely social forces. We exploit rotational symmetries and conservation laws of the problem to formulate a theory in terms of generalized coordinates of motion for the velocity directions akin to a Hamiltonian formulation for rotations. We explicitly derive the correspondence between this formulation and the dynamics of the individual velocities, thus obtaining a new model of collective motion. In the appropriate overdamped limit we recover the well-known Vicsek model, which dissipates rotational information and does not allow for polarized turns. Although the new model has its most vivid success in describing turning groups, its dynamics is intrinsically different from previous ones in a wide dynamical regime, while reducing to the hydrodynamic description of Toner and Tu at very large length-scales. The derived framework is therefore general and it may describe the collective motion of any strongly polarized active matter system.

GReTA-A Novel Global and Recursive Tracking Algorithm in Three Dimensions

Tracking multiple moving targets allows quantitative measure of the dynamic behavior in systems as diverse as animal groups in biology, turbulence in fluid dynamics and crowd and traffic control. In three dimensions, tracking several targets becomes increasingly hard since optical occlusions are very likely, i.e., two featureless targets frequently overlap for several frames. Occlusions are particularly frequent in biological groups such as bird flocks, fish schools, and insect swarms, a fact that has severely limited collective animal behavior field studies in the past. This paper presents a 3D tracking method that is robust in the case of severe occlusions. To ensure robustness, we adopt a global optimization approach that works on all objects and frames at once. To achieve practicality and scalability, we employ a divide and conquer formulation, thanks to which the computational complexity of the problem is reduced by orders of magnitude. We tested our algorithm with synthetic data, with experimental data of bird flocks and insect swarms and with public benchmark datasets, and show that our system yields high quality trajectories for hundreds of moving targets with severe overlap. The results obtained on very heterogeneous data show the potential applicability of our method to the most diverse experimental situations.

Nonsymmetric Interactions Trigger Collective Swings in Globally Ordered Systems

Many systems in nature, from ferromagnets to flocks of birds, exhibit ordering phenomena on the large scale. In condensed matter systems, order is statistically robust for large enough dimensions, with relative fluctuations due to noise vanishing with system size. Several biological systems, however, are less stable and spontaneously change their global state on relatively short time scales. Here we show that there are two crucial ingredients in these systems that enhance the effect of noise, leading to collective changes of state on finite time scales and off-equilibrium behavior: the nonsymmetric nature of interactions between individuals, and the presence of local heterogeneities in the topology of the network. Our results might explain what is observed in several living systems and are consistent with recent experimental data on bird flocks and other animal groups.