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Abstract The coordinated motion of animal groups through fluids is thought to reduce the cost of locomotion to individuals in the group. However, the connection between the spatial patterns observed in collectively moving animals and the energetic benefits at each position within the group remains unclear. To address this knowledge gap, we study the spontaneous emergence of cohesive formations in groups of fish, modeled as flapping foils, all heading in the same direction. We show in pairwise formations and with increasing group size that (1) in side-by-side arrangements, the reciprocal nature of flow coupling results in an equal distribution of energy re-quirements among all members, with reduction in cost of locomotion for swimmers flapping inphase but an increase in cost for swimmers flapping antiphase, and (2) in inline arrangements, flow coupling is non-reciprocal for all flapping phase, with energetic savings in favor of trailing swimmers, but only up to a finite number of swimmers, beyond which school cohesion and energetic benefits are lost at once. We explain these findings mechanistically and we provide efficient diagnostic tools for identifying locations in the wake of single and multiple swimmers that offer op-portunities for hydrodynamic benefits to aspiring followers. Our results imply a connection between the resources generated by flow physics and social traits that influence greedy and cooperative group behavior. 

The collective patterns that emerge in schooling fish are often analyzed using models of self-propelled particles in unbounded domains. However, while schooling fish in both field and laboratory settings interact with domain boundaries, these effects are typically ignored. Here, we propose a model that incorporates geometric confinement, by accounting for both flow and wall interactions, into existing data-driven behavioral rules. We show that new collective phases emerge where the school of fish “follows the tank wall” or “double mills.” Importantly, confinement induces repeated switching between two collective states, schooling and milling. We describe the group dynamics probabilistically, uncovering bistable collective states along with unintuitive bifurcations driving phase transitions. Our findings support the hypothesis that collective transitions in fish schools could occur spontaneously, with no adjustment at the individual level, and opens venues to control and engineer emergent collective patterns in biological and synthetic systems that operate far from equilibrium. 

We present experiments on oscillating hydrofoils undergoing combined heaving and pitching motions, paying particular attention to connections between propulsive efficiency and coherent wake features extracted using modal analysis. Time-averaged forces and particle image velocimetry measurements of the flow field downstream of the foil are presented for a Reynolds number of Re=11000 and Strouhal numbers in the range St=0.16--0.35. These conditions produce 2S and 2P wake patterns, as well as a near-momentumless wake structure. A triple decomposition using the optimized dynamic mode decomposition method is employed to identify dominant modal components (or coherent structures) in the wake. These structures can be connected to wake instabilities predicted using spatial stability analyses. Examining the modal components of the wake provides insightful explanations into the transition from drag to thrust production, and conditions that lead to peak propulsive efficiency. In particular, we find modes that correspond to the primary vortex development in the wakes. Other modal components capture elements of bluff body shedding at Strouhal numbers below the optimum for peak propulsive efficiency and characteristics of separation for Strouhal numbers higher than the optimum. 

A hydrostatic skeleton allows a soft body to transmit muscular force via internal pressure. A human's tongue, an octopus' arm and a nematode's body illustrate the pervasive presence of hydrostatic skeletons among animals, which has inspired the design of soft engineered actuators. However, there is a need for a theoretical basis for understanding how hydrostatic skeletons apply mechanical work. We therefore modeled the shape change and mechanics of natural and engineered hydrostatic skeletons to determine their mechanical advantage (MA) and displacement advantage (DA). These models apply to a variety of biological structures, but we explicitly consider the tube feet of a sea star and the body segments of an earthworm, and contrast them with a hydraulic press and a McKibben actuator. A helical winding of stiff, elastic fibers around these soft actuators plays a critical role in their mechanics by maintaining a cylindrical shape, distributing forces throughout the structure and storing elastic energy. In contrast to a single-joint lever system, soft hydrostats exhibit variable gearing with changes in MA generated by deformation in the skeleton. We found that this gearing is affected by the transmission efficiency of mechanical work (MA×DA) or, equivalently, the ratio of output to input work. The transmission efficiency changes with the capacity to store elastic energy within helically wrapped fibers or associated musculature. This modeling offers a conceptual basis for understanding the relationship between the morphology of hydrostatic skeletons and their mechanical performance. 

Fish typically swim by periodic bending of their bodies. Bending seems to follow a universal rule; it occurs at about one-third from the posterior end of the fish body with a maximum bending angle of about $$30^{\circ }$$ . However, the hydrodynamic mechanisms that shaped this convergent design and its potential benefit to fish in terms of swimming speed and efficiency are not well understood. It is also unclear to what extent this bending is active or follows passively from the interaction of a flexible posterior with the fluid environment. Here, we use a self-propelled two-link model, with fluid–structure interactions described in the context of the vortex sheet method, to analyse the effects of both active and passive body bending on the swimming performance. We find that passive bending is more efficient but could reduce swimming speed compared with rigid flapping, but the addition of active bending could enhance both speed and efficiency. Importantly, we find that the phase difference between the posterior and anterior sections of the body is an important kinematic factor that influences performance, and that active antiphase flexion, consistent with the passive flexion phase, can simultaneously enhance speed and efficiency in a region of the design space that overlaps with biological observations. Our results are consistent with the hypothesis that fish that actively bend their bodies in a fashion that exploits passive hydrodynamics can at once improve speed and efficiency. 

Fish schools are ubiquitous in marine life. Although flow interactions are thought to be beneficial for schooling, their exact effects on the speed, energetics and stability of the group remain elusive. Recent numerical simulations and experimental models suggest that flow interactions stabilize in-tandem formations of flapping foils. Here, we employ a minimal vortex sheet model that captures salient features of the flow interactions among flapping swimmers, and we study the free swimming of a pair of in-line swimmers driven with identical heaving or pitching motions. We find that, independent of the flapping mode, heaving or pitching, the follower passively stabilizes at discrete locations in the wake of the leader, consistent with the heaving foil experiments, but pitching swimmers exhibit tighter and more cohesive formations. Further, in comparison to swimming alone, pitching motions increase the energetic efficiency of the group while heaving motions result in a slight increase in the swimming speed. A deeper analysis of the wake of a single swimmer sheds light on the hydrodynamic mechanisms underlying pairwise formations. These results recapitulate that flow interactions provide a passive mechanism that promotes school cohesion, and afford novel insights into the role of the flapping mode in controlling the emergent properties of the school.