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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.more » « lessFree, publicly-accessible full text available February 15, 2026
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            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.more » « lessFree, publicly-accessible full text available December 1, 2025
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            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.more » « less
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            ABSTRACT Humans have been trying to understand animal behavior at least since recorded history. Recent rapid development of new technologies has allowed us to make significant progress in understanding the physiological and molecular mechanisms underlying behavior, a key goal of neuroethology. However, there is a tradeoff when studying animal behavior and its underlying biological mechanisms: common behavior protocols in the laboratory are designed to be replicable and controlled, but they often fail to encompass the variability and breadth of natural behavior. This Commentary proposes a framework of 10 key questions that aim to guide researchers in incorporating a rich natural context into their experimental design or in choosing a new animal study system. The 10 questions cover overarching experimental considerations that can provide a template for interspecies comparisons, enable us to develop studies in new model organisms and unlock new experiments in our quest to understand behavior.more » « less
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            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.more » « less
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            Abstract Many aquatic animals swim by undulatory body movements and understanding the diversity of these movements could unlock the potential for designing better underwater robots. Here, we analyzed the steady swimming kinematics of a diverse group of fish species to investigate whether their undulatory movements can be represented using a series of interconnected multi-segment models, and if so, to identify the key factors driving the segment configuration of the models. Our results show that the steady swimming kinematics of fishes can be described successfully using parsimonious models, 83% of which had fewer than five segments. In these models, the anterior segments were significantly longer than the posterior segments, and there was a direct link between segment configuration and swimming kinematics, body shape, and Reynolds number. The models representing eel-like fishes with elongated bodies and fishes swimming at high Reynolds numbers had more segments and less segment length variability along the body than the models representing other fishes. These fishes recruited their anterior bodies to a greater extent, initiating the undulatory wave more anteriorly. Two shape parameters, related to axial and overall body thickness, predicted segment configuration with moderate to high success rate. We found that head morphology was a good predictor of its segment length. While there was a large variation in head segments, the length of tail segments was similar across all models. Given that fishes exhibited variable caudal fin shapes, the consistency of tail segments could be a result of an evolutionary constraint tuned for high propulsive efficiency. The bio-inspired multi-segment models presented in this study highlight the key bending points along the body and can be used to decide on the placement of actuators in fish-inspired robots, to model hydrodynamic forces in theoretical and computational studies, or for predicting muscle activation patterns during swimming.more » « less
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            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.more » « lessFree, publicly-accessible full text available January 1, 2026
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            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.more » « less
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            Animals can evolve dramatic sensory functions in response to environmental constraints, but little is known about the neural mechanisms underlying these changes. The Mexican tetra, Astyanax mexicanus , is a leading model to study genetic, behavioral, and physiological evolution by comparing eyed surface populations and blind cave populations. We compared neurophysiological responses of posterior lateral line afferent neurons and motor neurons across A. mexicanus populations to reveal how shifts in sensory function may shape behavioral diversity. These studies indicate differences in intrinsic afferent signaling and gain control across populations. Elevated endogenous afferent activity identified a lower response threshold in the lateral line of blind cavefish relative to surface fish leading to increased evoked potentials during hair cell deflection in cavefish. We next measured the effect of inhibitory corollary discharges from hindbrain efferent neurons onto afferents during locomotion. We discovered that three independently derived cavefish populations have evolved persistent afferent activity during locomotion, suggesting for the first time that partial loss of function in the efferent system can be an evolutionary mechanism for neural adaptation of a vertebrate sensory system.more » « less
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