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ABSTRACT Most of our understanding of fish locomotion has focused on elementary behaviors such as steady swimming and escape responses in simple environments. As the field matures, increasing attention is being paid to transient and unsteady behaviors that characterize more complex interactions with the environment. This Commentary advocates for an ecologically relevant approach to lab studies. Specific examples have brought new understanding to the energetic consequences of fish swimming, such as (1) station holding around bluff bodies, which departs drastically from steady swimming in almost all aspects of kinematics, muscle activity and energetics, and (2) transient behaviors such as acceleration and feeding, which are critical to survival but often neglected because of challenges in measuring costs. Beyond the lab, a far richer diversity of behaviors is available when fish are given enough space and time to move. Mesocosm studies are poised to reveal new insights into fish swimming that are inaccessible in laboratory settings. Next-generation biologgers that incorporate neural recordings will usher in a new era for understanding biomechanics in the wild and open the door for a more mechanistic understanding of how changing environments affect animal movement. These advances promise to allow insights into animal locomotion in ways that will mutually complement and accelerate laboratory and field studies in the years to come. 

ABSTRACT Fish in the wild often contend with complex flows that are produced by natural and artificial structures. Research into fish interactions with turbulence often investigates metrics such as turbulent kinetic energy (TKE) or fish positional location, with less focus on the specific interactions between vortex organization and body swimming kinematics. Here, we compared the swimming kinematics of rainbow trout (Oncorhynchus mykiss) holding station in flows produced by two different 3×5 cylinder arrays. We systematically utilized computational fluid dynamics to identify one array that produced a Kármán vortex street with high vortex periodicity (KVS array) and another that produced low periodicity, similar to a parallel vortex street (PVS array), both validated with particle image velocimetry. The only difference in swimming kinematics between cylinder arrays was an increased tail beat amplitude in the KVS array. In both cylinder arrays, the tail beat frequency decreased and snout amplitude increased compared with the freestream. The center of mass amplitude was greater in the PVS array than in only the freestream, however, suggesting some buffeting of the body by the fluid. Notably, we did not observe Kármán gaiting in the KVS array as in previous studies. We hypothesize that this is because (1) vorticity was dissipated in the region where fish held station or (2) vortices were in-line rather than staggered. These results are the first to quantify the kinematics and behavior of fishes swimming in the wake of multiple cylinder arrays, which has important implications for biomechanics, fluid dynamics and fisheries management. 

ABSTRACT Complex hydrodynamics abound in natural streams, yet the selective pressures these impose upon different size classes of fish are not well understood. Attached vortices are produced by relatively large objects that block freestream flow, which fish routinely utilize for flow refuging. To test how flow refuging and the potential harvesting of energy (as seen in Kármán gaiting) vary across size classes in rainbow trout (Oncorhynchus mykiss; fingerling, 8 cm; parr, 14 cm; adult, 22 cm; n=4 per size class), we used a water flume (4100 l; freestream flow at 65 cm s−1) and created vortices using 45 deg wing dams of varying size (small, 15 cm; medium, 31 cm; large, 48 cm). We monitored microhabitat selection and swimming kinematics of individual trout and measured the flow field in the wake of wing dams using time-resolved particle image velocimetry (PIV). Trout of each size class preferentially swam in vortices rather than the freestream, but the capacity to flow refuge varied according to the ratio of vortex width to fish length (WV:LF). Consistent refuging behavior was exhibited when WV:LF≥1.5. All size classes exhibited increased wavelength and Strouhal number and decreased tailbeat frequency within vortices compared with freestream, suggesting that swimming in vortices requires less power output. In 17% of the trials, fish preferentially swam in a manner that suggests energy harvesting from the shear layer. Our results can inform efforts toward riparian restoration and fishway design to improve salmonid conservation. 

ABSTRACT Pectoral fins play a crucial role in fish locomotion. Despite fishes living in complex fluid environments that exist in rivers and tidal flows, the role of the pectoral fins in navigating turbulent flows is not well understood. This study investigated the kinematics and muscle activity of pectoral fins in rainbow trout as they held station in the unsteady flows behind a D-section cylinder. We observed two distinct pectoral fin behaviors, one during braking and the other during Kármán gaiting. These behaviors were correlated to whole-body movements in response to the hydrodynamic conditions of specific regions in the cylinder wake. Sustained fin extensions during braking, where the fin was held out to maintain its position away from the body and against the flow, were associated with the cessation of forward body velocity, where the fish avoided the suction region directly downstream of the cylinder. Transient fin extensions and retractions during Kármán gaiting controlled body movements in the cross-stream direction. These two fin behaviors had different patterns of muscle activity. All braking events required recruitment from both the abductor and adductor musculature to actively extend a pectoral fin. In contrast, over 50% of fin extension movements during Kármán gaiting proceed in the absence of muscle activity. We reveal that in unsteady fluid environments, pectoral fin movements are the result of a complex combination of passive and active mechanisms that deviate substantially from canonical labriform locomotion, the implications of which await further work on the integration of sensory and motor systems. 

Mobulid rays (manta and devil rays) use a highly specialized filtering apparatus to separate plankton food particles from seawater. Recent studies have indicated that captive vortices form within the microscale pores of the filter, which enhance filtration efficiency through a novel mechanism referred to as ricochet separation. The high throughput and clog resistance of this filtration process have led to the development of several bioinspired engineered filtration systems. However, it is still unclear how changes to the filter morphology influence the surrounding flow patterns and filtration efficiency. We address this question by examining the flow fields around and filtering properties of mobulid filters with systematically varied morphologies, using a combination of computational fluid dynamics and experiments on physical models. While the pore size is the principal determinant of filtration efficiency in a sieve filter, we found that the captive vortices in a mobulid filter grow or shrink to fill the pore, and changes in the pore size have modest effects. By contrast, the filtration efficiency appears to be highly sensitive to the orientation of the filter lobes (microscale plate-like structures). These results provide a foundation for interpreting the morphological differences between species and also for generating optimized bioinspired designs. 

Predator–prey interactions are fundamental to ecological and evolutionary dynamics. Yet, predicting the outcome of such interactions—whether predators intercept prey or fail to do so—remains a challenge. An emerging hypothesis holds that interception trajectories of diverse predator species can be described by simple feedback control laws that map sensory inputs to motor outputs. This form of feedback control is widely used in engineered systems but suffers from degraded performance in the presence of processing delays such as those found in biological brains. We tested whether delay-uncompensated feedback control could explain predator pursuit manoeuvres using a novel experimental system to present hunting fish with virtual targets that manoeuvred in ways that push the limits of this type of control. We found that predator behaviour cannot be explained by delay-uncompensated feedback control, but is instead consistent with a pursuit algorithm that combines short-term forecasting of self-motion and prey motion with feedback control. This model predicts both predator interception trajectories and whether predators capture or fail to capture prey on a trial-by-trial basis. Our results demonstrate how animals can combine short-term forecasting with feedback control to generate robust flexible behaviours in the face of significant processing delays.