Search

Hydrostatic skeletons, such as an elephant trunk or a squid tentacle, permit the transmission of mechanical work through a soft body. Despite the ubiquity of these structures among animals, we generally do not understand how differences in their morphology affect their ability to transmit muscular work. Therefore, the present study used mathematical modeling, morphometrics, and kinematics to understand the transmission of force and displacement in the tube feet of the juvenile six-rayed star (Leptasterias sp.). An inverse-dynamic analysis revealed that the forces generated by the feet during crawling primarily serve to overcome the submerged weight of the body. These forces were disproportionately generated by the feet at more proximal positions along each ray, which were used more frequently for crawling. Owing to a combination of mechanical advantage and muscle mass, these proximal feet exhibited a greater capacity for force generation than the distal feet. However, the higher displacement advantage of the more elongated distal feet offer a superior ability to extend the feet into the environment. Therefore, the morphology of tube feet demonstrates a gradient in gearing along each ray that compliments their role in behavior., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Schooling fish rely on a social network created through signaling between its members to interact with their environment. Previous studies have established that vision is necessary for schooling and that flow sensing by the lateral line system may aid in a school's cohesion. However, it remains unclear to what extent flow provides a channel of communication between schooling fish. Based on kinematic measurements of the speed and heading of schooling tetras (Petitella rhodostoma), we found that compromising the lateral line by chemical treatment reduced the mutual information between individuals by ∼13%. This relatively small reduction in pairwise communication propagated through schools of varying size to reduce the degree and connectivity of the social network by more than half. Treated schools additionally showed more than twice the spatial heterogeneity of fish with unaltered flow sensing. These effects were much more substantial than the changes that we measured in the nearest-neighbor distance, speed and intermittency of individual fish by compromising flow sensing. Therefore, flow serves as a valuable supplement to visual communication in a manner that is revealed through a school's network properties., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

It is unclear how animals with radial symmetry control locomotion without a brain. Using a combination of experiments, mathematical modeling, and robotics, we tested the extent to which this control emerges in sea stars (Protoreaster nodosus) from the local control of their hundreds of feet and their mechanical interactions with the body. We discovered that these animals compensate for an experimental increase in their submerged weight by recruiting more feet that synchronize in the power stroke of the locomotor cycle during their bouncing gait. Mathematical modeling of the mechanics of a sea star replicated this response to loading without a central controller. A robotic sea star was found to similarly recruit more actuators under higher loads through purely decentralized control. These results suggest that an array of biological or engineered actuators are capable of cooperative transport where the actuators are dynamically recruited by the mechanics of the body. In particular, the body's vertical oscillations serve to recruit feet in greater numbers to overcome the weight to propel the body forward. This form of distributed control contrasts the conventional view of animal locomotion as governed by the central nervous system and offers inspiration for the design of engineered devices with arrays of actuators., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 Elsevier Inc. All rights reserved.)

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., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

To celebrate its centenary year, Journal of Experimental Biology (JEB) commissioned a collection of articles examining the past, present and future of experimental biology. This Commentary closes the collection by considering the important research opportunities and challenges that await us in the future. We expect that researchers will harness the power of technological advances, such as '-omics' and gene editing, to probe resistance and resilience to environmental change as well as other organismal responses. The capacity to handle large data sets will allow high-resolution data to be collected for individual animals and to understand population, species and community responses. The availability of large data sets will also place greater emphasis on approaches such as modeling and simulations. Finally, the increasing sophistication of biologgers will allow more comprehensive data to be collected for individual animals in the wild. Collectively, these approaches will provide an unprecedented understanding of 'how animals work' as well as keys to safeguarding animals at a time when anthropogenic activities are degrading the natural environment., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

This paper investigates a pursuit-evasion game with a single pursuer and evader in a bounded environment, inspired by observations of predation attempts by lionfish ( Pterois sp. ). The pursuer tracks the evader with a pure pursuit strategy while using an additional bioinspired tactic to trap the evader, i.e. minimize the evader's escape routes. Specifically, the pursuer employs symmetric appendages inspired by the large pectoral fins of lionfish, but this expansion increases its drag and therefore its work to capture the evader. The evader employs a bioinspired randomly-directed escape strategy to avoid capture and collisions with the boundary. Here we investigate the trade-off between minimizing the work to capture the evader and minimizing the evader's escape routes. By using the pursuer's expected work to capture as a cost function, we determine when the pursuer should expand its appendages as a function of the relative distance to the evader and the evader's proximity to the boundary. Visualizing the pursuer's expected work to capture everywhere in the bounded domain, yields additional insights about optimal pursuit trajectories and illustrates the role of the boundary in predator-prey interactions., (Creative Commons Attribution license.)

An effective evasion strategy allows prey to survive encounters with predators. Prey are generally thought to escape in a direction that is either random or serves to maximize the minimum distance from the predator. Here, we introduce a comprehensive approach to determine the most likely evasion strategy among multiple hypotheses and the role of biomechanical constraints on the escape response of prey fish. Through a consideration of six strategies with sensorimotor noise and previous kinematic measurements, our analysis shows that zebrafish larvae generally escape in a direction orthogonal to the predator's heading. By sensing only the predator's heading, this orthogonal strategy maximizes the distance from fast-moving predators, and, when operating within the biomechanical constraints of the escape response, it provides the best predictions of prey behavior among all alternatives. This work demonstrates a framework for resolving the strategic basis of evasion in predator-prey interactions, which could be applied to a broad diversity of animals.

Kinematic measurements have been essential to the study of comparative biomechanics and offer insight into relationships between technological development and scientific progress. Here, we review the 100 year history of kinematic measurements in Journal of Experimental Biology (JEB) through eras that used film, analog video and digital video, and approaches that have circumvented the use of image capture. This history originated with the career of Sir James Gray and has since evolved over the generations of investigators that have followed. Although some JEB studies have featured technological developments that were ahead of their time, the vast majority of research adopted equipment that was broadly available through the consumer or industrial markets. We found that across eras, an emphasis on high-speed phenomena outpaced the growth of the number of articles published by JEB and the size of datasets increased significantly. Despite these advances, the number of species studied within individual reports has not differed significantly over time. Therefore, we find that advances in technology have helped to enable a growth in the number of JEB studies that have included kinematic measurements, contributed to an emphasis on high-speed phenomena, and yielded biomechanical studies that are more data rich, but are no more comparative now than in previous decades., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

The pursuit of prey is vital to the biology of a predator and many aspects of predatory behaviour are well-studied. However, it is unclear how a pursuit can be effective when the prey is faster than a non-cryptic predator. Using kinematic measurements, we considered the strategy of red lionfish ( Pterois volitans ) as they pursued a faster prey fish ( Chromis viridis ) under laboratory conditions. Despite swimming about half as fast as C. viridis , lionfish succeeded in capturing prey in 61% of our experiments. This successful pursuit behaviour was defined by three critical characteristics. First, lionfish targeted C. viridis with pure pursuit by adjusting their heading towards the prey's position and not the anticipated point of interception. Second, lionfish pursued prey with uninterrupted motion. By contrast, C. viridis moved intermittently with variation in speed that included slow swimming. Such periods allowed lionfish to close the distance to a prey and initiate a suction-feeding strike at a relatively close distance (less than 9 cm). Finally, lionfish exhibited a high rate of strike success, capturing prey in 74% of all strikes. These characteristics comprise a behaviour that we call the 'persistent-predation strategy', which may be exhibited by a diversity of predators with relatively slow locomotion.

Predator-prey interactions are critical to the biology of a diversity of animals. Although prey capture is determined by the direction, velocity, and timing of motion by both animals, it is generally unclear what strategies are employed by predators and prey to guide locomotion. Here we review our research on fishes that tests the pursuit strategy of predators and the evasion strategy of prey through kinematic measurements and agent-based models. This work demonstrates that fish predators track prey with variations on a deviated-pursuit strategy that is guided by visual cues. Fish prey employ a mixed strategy that varies with factors such as the direction of a predator's approach. Our models consider the stochastic nature of interactions by incorporating measured probability distributions to accurately predict measurements of survivorship. A sensitivity analysis of these models shows the importance of the response distance of prey to their survival. Collectively, this work demonstrates how strategy affects the outcome of predator-prey interactions and articulates the roles of sensing, control, and propulsion. The research program that we have developed has the potential to offer a framework for the study of strategy in the predator-prey interactions of a variety of animals., (© The Author(s) 2021. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

Research that integrates animal behavior theory with mechanics-including biomechanics, physiology, and functional morphology-can reveal how organisms accomplish tasks crucial to their fitness. Despite the insights that can be gained from this interdisciplinary approach, biomechanics commonly neglects a behavioral context and behavioral research generally does not consider mechanics. Here, we aim to encourage the study of "mechanoethology," an area of investigation intended to encompass integrative studies of mechanics and behavior. Using examples from the literature, including papers in this issue, we show how these fields can influence each other in three ways: (1) the energy required to execute behaviors is driven by the kinematics of movement, and mechanistic studies of movement can benefit from consideration of its behavioral context; (2) mechanics sets physical limits on what behaviors organisms execute, while behavior influences ecological and evolutionary limits on mechanical systems; and (3) sensory behavior is underlain by the mechanics of sensory structures, and sensory systems guide whole-organism movement. These core concepts offer a foundation for mechanoethology research. However, future studies focused on merging behavior and mechanics may reveal other ways by which these fields are linked, leading to further insights in integrative organismal biology., (© The Author(s) 2021. Published by Oxford University Press on behalf of Integrative and Comparative Biology.)

Latch-mediated spring actuation (LaMSA) is used by small organisms to produce high acceleration movements. Mathematical models predict that acceleration increases as LaMSA systems decrease in size. Adult mantis shrimp use a LaMSA mechanism in their raptorial appendages to produce extremely fast strikes. Until now, however, it was unclear whether mantis shrimp at earlier life-history stages also strike using elastic recoil and latch mediation. We tested whether larval mantis shrimp (Gonodactylaceus falcatus) use LaMSA and, because of their smaller size, achieve higher strike accelerations than adults of other mantis shrimp species. Based on microscopy and kinematic analyses, we discovered that larval G. falcatus possess the components of, and actively use, LaMSA during their fourth larval stage, which is the stage of development when larvae begin feeding. Larvae performed strikes at high acceleration and speed (mean: 4.133×105 rad s-2, 292.7 rad s-1; 12 individuals, 25 strikes), which are of the same order of magnitude as for adults - even though adult appendages are up to two orders of magnitude longer. Larval strike speed (mean: 0.385 m s-1) exceeded the maximum swimming speed of similarly sized organisms from other species by several orders of magnitude. These findings establish the developmental timing and scaling of the mantis shrimp LaMSA mechanism and provide insights into the kinematic consequences of scaling limits in tiny elastic mechanisms., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

The control of a predator's locomotion is critical to its ability to capture prey. Flying animals adjust their heading continuously with control similar to guided missiles. However, many animals do not move with rapid continuous motion, but rather interrupt their progress with frequent pauses. To understand how such intermittent locomotion may be controlled during predation, we examined the kinematics of zebrafish ( Danio rerio ) as they pursued larval prey of the same species. Like many fishes, zebrafish move with discrete burst-and-coast swimming. We found that the change in heading and tail excursion during the burst phase was linearly related to the prey's bearing. These results suggest a strategy, which we call intermittent pure pursuit, that offers advantages in sensing and control. This control strategy is similar to perception and path-planning algorithms required in the design of some autonomous robots and may be common to a diversity of animals., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Schooling is a collective behaviour that enhances the ability of a fish to sense and respond to its environment. Although schooling is essential to the biology of a diversity of fishes, it is generally unclear how this behaviour is coordinated by different sensory modalities. We used experimental manipulation and kinematic measurements to test the role of vision and flow sensing in the rummy-nose tetra ( Hemigrammus rhodostomus ), which swims with intermittent phases of bursts and coasts. Groups of five fish required a minimum level of illuminance (greater than 1.5 lx) to achieve the necessary close nearest-neighbour distance and high polarization for schooling. Compromising the lateral line system with an antibiotic treatment caused tetras to swim with greater nearest-neighbour distance and lower polarization. Therefore, vision is both necessary and sufficient for schooling in H. rhodostomus , and both sensory modalities aid in attraction. These results can serve as a basis for understanding the individual roles of sensory modalities in schooling for some fish species.

A diversity of animals survive encounters with predators by escaping from a looming visual stimulus. Despite the importance of this behavior, it is generally unclear how visual cues facilitate a prey's survival from predation. Therefore, the aim of this study was to understand how the visual angle subtended on the eye of the prey by the predator affects the distance of adult zebrafish ( Danio rerio ) from predators. We performed experiments to measure the threshold visual angle and mathematically modeled the kinematics of predator and prey. We analyzed the responses to the artificial stimulus with a novel approach that calculated relationships between hypothetical values for a threshold-stimulus angle and the latency between stimulus and response. These relationships were verified against the kinematic responses of zebrafish to a live fish predator ( Herichthys cyanoguttatus ). The predictions of our model suggest that the measured threshold visual angle facilitates escape when the predator's approach is slower than approximately twice the prey's escape speed. These results demonstrate the capacity and limits to how the visual angle provides a prey with the means to escape a predator., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

The oral surface of sea stars is lined with arrays of tube feet that enable them to achieve highly controlled locomotion on various terrains. The activity of the tube feet is orchestrated by a nervous system that is distributed throughout the body without a central brain. How such a distributed nervous system produces a coordinated locomotion is yet to be understood. We develop mathematical models of the biomechanics of the tube feet and the sea star body. In the model, the feet are coupled mechanically through their structural connection to a rigid body. We formulate hierarchical control laws that capture salient features of the sea star nervous system. Namely, at the tube foot level, the power and recovery strokes follow a state-dependent feedback controller. At the system level, a directionality command is communicated through the nervous system to all tube feet. We study the locomotion gaits afforded by this hierarchical control model. We find that these minimally coupled tube feet coordinate to generate robust forward locomotion, reminiscent of the crawling motion of sea stars, on various terrains and for heterogeneous tube feet parameters and initial conditions. Our model also predicts a transition from crawling to bouncing consistently with recent experiments. We conclude by commenting on the implications of these findings for understanding the neuromechanics of sea stars and their potential application to autonomous robotic systems.

Biologists commonly visualize different features of an organism using distinct sources of illumination. Such multichannel imaging has largely not been applied to behavioral studies because of the challenges posed by a moving subject. We address this challenge with the technique of multichannel stroboscopic videography (MSV), which synchronizes multiple strobe lights with video exposures of a single camera. We illustrate the utility of this approach with kinematic measurements of a walking cockroach ( Gromphadorhina portentosa ) and calculations of the pressure field around a swimming fish ( Danio rerio ). In both, transmitted illumination generated high-contrast images of the animal's body in one channel. Other sources of illumination were used to visualize the points of contact for the feet of the cockroach and the water flow around the fish in separate channels. MSV provides an enhanced potential for high-throughput experimentation and the capacity to integrate changes in physiological or environmental conditions in freely-behaving animals., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Aquatic animals commonly sense flow using superficial neuromasts (SNs), which are receptors that extend from the body's surface. The lateral line of fishes is unique among these systems because it additionally possesses receptors, the canal neuromasts (CNs), that are recessed within a channel. The lateral line has inspired the development of engineered sensors and concepts in the analysis of flow fields for submersible navigation. The biophysics of CNs are known to be different from the SNs and thereby offer a distinct submodality. However, it is generally unclear whether CNs play a distinct role in behavior. We therefore tested whether CNs enhance foraging in the dark by zebrafish (Danio rerio), a behavior that we elicited with a vibrating rod. We found that juvenile fish, which have only SNs, bite at this rod at about one-third the rate and from as little as one-third the distance of adults for a high-frequency stimulus (50  <  f   <  100 Hz). We used novel techniques for manipulating the lateral line in adults to find that CNs offered only a modest benefit at a lower frequency (20 Hz) and that foraging was mediated entirely by cranial neuromasts. Consistent with our behavioral results, biophysical models predicted CNs to be more than an order of magnitude more sensitive than SNs at high frequencies. This enhancement helps to overcome the rapid spatial decay in high-frequency components in the flow around the stimulus. These findings contrast what has been previously established for fishes that are at least ten-times the length of zebrafish, which use trunk CNs to localize prey. Therefore, CNs generally enhance foraging, but in a manner that varies with the size of the fish and its prey. These results have the potential to improve our understanding of flow sensing in aquatic animals and engineered systems.

Measurements of energy use, and its scaling with size, are critical to understanding how organisms accomplish myriad tasks. For example, energy budgets are central to game theory models of assessment during contests and underlie patterns of feeding behavior. Clear tests connecting energy to behavioral theory require measurements of the energy use of single individuals for particular behaviors. Many species of mantis shrimp (Stomatopoda: Crustacea) use elastic energy storage to power high-speed strikes that they deliver to opponents during territorial contests and to hard-shelled prey while feeding. We compared the scaling of strike kinematics and energetics between feeding and contests in the mantis shrimp Neogonodactylus bredini We filmed strikes with high-speed video, measured strike velocity and used a mathematical model to calculate strike energy. During contests, strike velocity did not scale with body size but strike energy scaled positively with size. Conversely, while feeding, strike velocity decreased with increasing size and strike energy did not vary according to body size. Individuals most likely achieved this strike variation through differential compression of their exoskeletal spring prior to the strike. Post hoc analyses found that N. bredini used greater velocity and energy when striking larger opponents, yet variation in prey size was not accompanied by varying strike velocity or energetics. Our estimates of energetics inform prior tests of contest and feeding behavior in this species. More broadly, our findings elucidate the role behavioral context plays in measurements of animal performance., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

A predator's ability to capture prey depends critically on how it coordinates its approach in response to a prey's motion. Flying insects, bats and raptors are capable of capturing prey with a strategy known as parallel navigation, which allows a predator to move directly towards the anticipated point of interception. It is unclear if predators using other modes of locomotion are employing this strategy when pursuing evasive prey. Using kinematic measurements and mathematical modelling, we tested whether bluefish ( Pomatomus saltatrix) pursue prey fish ( Fundulus heteroclitus) with parallel navigation. We found that the directional changes of bluefish were not consistent with this strategy, but rather were predicted by a strategy known as deviated pursuit. Although deviated pursuit requires few sensory cues and relatively modest motor coordination, a comparison of mathematical models suggested negligible differences in path length from parallel navigation, largely owing to the acceleration exhibited by bluefish near the end of a pursuit. Therefore, the strategy of bluefish is unlike flying predators, but offers comparable performance with potentially more robust control that may be well suited to the visual system and habitat of fishes. These findings offer a foundation for understanding the sensing and locomotor control of predatory fishes.

Predation is a fundamental interaction between species, yet it is largely unclear what tactics are successful for the survival or capture of prey. One challenge in this area comes with how to test theoretical ideas about strategy with experimental measurements of features such as speed, flush distance and escape angles. Tactics may be articulated with an analytical model that predicts the motion of predator or prey as they interact. However, it may be difficult to recognize how the predictions of such models relate to behavioural measurements that are inherently variable. Here, we present an alternative approach for modelling predator-prey interactions that uses deterministic dynamics, yet incorporates experimental kinematic measurements of natural variation to predict the outcome of biological events. This technique, called probabilistic analytical modelling (PAM), is illustrated by the interactions between predator and prey fish in two case studies that draw on recent experiments. In the first case, we use PAM to model the tactics of predatory bluefish ( Pomatomus saltatrix) as they prey upon smaller fish ( Fundulus heteroclitus). We find that bluefish perform deviated pure pursuit with a variable pursuit angle that is suboptimal for the time to capture. In the second case, we model the escape tactics of zebrafish larvae ( Danio rerio) when approached by adult predators of the same species. Our model successfully predicts the measured patterns of survivorship using measured probability density functions as parameters. As these results demonstrate, PAM is a data-driven modelling approach that can be predictive, offers analytical transparency, and does not require numerical simulations of system dynamics. Though predator-prey interactions demonstrate the use of this technique, PAM is not limited to studying biological systems and has broad utility that may be applied towards understanding a wide variety of natural and engineered dynamical systems where data-driven modelling is beneficial.

Predation is a fundamental interaction between species, yet it is unclear what escape strategies are effective for prey survival. Classical theory proposes that prey should either escape in a direction that conforms to a performance optimum or that is random and therefore unpredictable. Here, we show that larval zebrafish ( Danio rerio ) instead use a mixed strategy that may be either random or directed. This was determined by testing classic theory with measurements of the escape direction in response to a predator robot. We found that prey consistently escaped in a direction contralateral to the robot when approached from the side of the prey's body. At such an orientation, the predator appeared in the prey's central visual field and the contralateral response was consistent with a model of strategy that maximizes the distance from the predator. By contrast, when the robot approached the rostral or caudal ends of the body, and appeared in the prey's peripheral vision, the escape showed an equal probability of a contralateral or ipsilateral direction. At this orientation, a contralateral response offered little strategic advantage. Therefore, zebrafish larvae adopt an escape strategy that maximizes distance from the threat when strategically beneficial and that is otherwise random. This sensory-mediated mixed strategy may be employed by a diversity of animals and offers a new paradigm for understanding the factors that govern prey survival., (© 2017 The Author(s).)

An escape response is a rapid manoeuvre used by prey to evade predators. Performing this manoeuvre at greater speed, in a favourable direction, or from a longer distance have been hypothesized to enhance the survival of prey, but these ideas are difficult to test experimentally. We examined how prey survival depends on escape kinematics through a novel combination of experimentation and mathematical modelling. This approach focused on zebrafish ( Danio rerio ) larvae under predation by adults and juveniles of the same species. High-speed three-dimensional kinematics were used to track the body position of prey and predator and to determine the probability of behavioural actions by both fish. These measurements provided the basis for an agent-based probabilistic model that simulated the trajectories of the animals. Predictions of survivorship by this model were found by Monte Carlo simulations to agree with our observations and we examined how these predictions varied by changing individual model parameters. Contrary to expectation, we found that survival may not be improved by increasing the speed or altering the direction of the escape. Rather, zebrafish larvae operate with sufficiently high locomotor performance due to the relatively slow approach and limited range of suction feeding by fish predators. We did find that survival was enhanced when prey responded from a greater distance. This is an ability that depends on the capacity of the visual and lateral line systems to detect a looming threat. Therefore, performance in sensing, and not locomotion, is decisive for improving the survival of larval fish prey. These results offer a framework for understanding the evolution of predator-prey strategy that may inform prey survival in a broad diversity of animals., (© 2017 The Author(s).)

Countless aquatic animals rotate appendages through the water, yet fluid forces are typically modeled with translational motion. To elucidate the hydrodynamics of rotation, we analyzed the raptorial appendages of mantis shrimp (Stomatopoda) using a combination of flume experiments, mathematical modeling and phylogenetic comparative analyses. We found that computationally efficient blade-element models offered an accurate first-order approximation of drag, when compared with a more elaborate computational fluid-dynamic model. Taking advantage of this efficiency, we compared the hydrodynamics of the raptorial appendage in different species, including a newly measured spearing species, Coronis scolopendra The ultrafast appendages of a smasher species (Odontodactylus scyllarus) were an order of magnitude smaller, yet experienced values of drag-induced torque similar to those of a spearing species (Lysiosquillina maculata). The dactyl, a stabbing segment that can be opened at the distal end of the appendage, generated substantial additional drag in the smasher, but not in the spearer, which uses the segment to capture evasive prey. Phylogenetic comparative analyses revealed that larger mantis shrimp species strike more slowly, regardless of whether they smash or spear their prey. In summary, drag was minimally affected by shape, whereas size, speed and dactyl orientation dominated and differentiated the hydrodynamic forces across species and sizes. This study demonstrates the utility of simple mathematical modeling for comparative analyses and illustrates the multi-faceted consequences of drag during the evolutionary diversification of rotating appendages., (© 2016. Published by The Company of Biologists Ltd.)

A large diversity of fishes struggle early in life to forage on zooplankton while under the threat of predation. Some species, such as zebrafish (Danio rerio), acquire an ability to forage in the dark during growth as larvae, but it is unclear how this is achieved. We investigated the functional basis of this foraging by video-recording larval and juvenile zebrafish as they preyed on zooplankton (Artemia sp.) under infrared illumination. We found that foraging improved with age, to the extent that 1-month-old juveniles exhibited a capture rate that was an order of magnitude greater than that of hatchlings. At all ages, the ability to forage in the dark was diminished when we used a chemical treatment to compromise the cranial superficial neuromasts, which facilitate flow sensing. However, a morphological analysis showed no developmental changes in these receptors that could enhance sensitivity. We tested whether the improvement in foraging with age could instead be a consequence of learning by raising fish that were naïve to the flow of prey. After 1 month of growth, both groups foraged with a capture rate that was significantly less than that of fish that had the opportunity to learn and indistinguishable from that of fish with no ability to sense flow. This suggests that larval fish learn to use water flow to forage in the dark. This ability could enhance resource acquisition under reduced competition and predation. Furthermore, our findings offer an example of learning in a model system that offers promise for understanding its neurophysiological basis., (© 2016. Published by The Company of Biologists Ltd.)

Larval fish use the 'fast start' escape response to rapidly evade the strike of a predator with a three-dimensional (3D) maneuver. Although this behavior is essential for the survival of fishes, it is not clear how its motion is controlled by the motor system of a larval fish. As a basis for understanding this control, we measured the high-speed kinematics of the body of zebrafish (Danio rerio) larvae when executing the fast start in a variety of directions. We found that the angular excursion in the lateral direction is correlated with the yaw angle in the initial stage of bending (stage 1). In this way, larvae moved in a manner similar to that reported for adult fish. However, larvae also have the ability to control the elevation of a fast start. We found that escapes directed downwards or upwards were achieved by pitching the body throughout the stages of the fast start. Changes in the pitching angle in each stage were significantly correlated with the elevation angle of the trajectory. Therefore, as a larva performs rapid oscillations in yaw that contribute to undulatory motion, the elevation of an escape is generated by more gradual and sustained changes in pitch. These observations are consistent with a model of motor control where elevation is directed through the differential activation of the epaxial and hypaxial musculature. This 3D motion could serve to enhance evasiveness by varying elevation without slowing the escape from a predator., (© 2015. Published by The Company of Biologists Ltd.)

Predator-prey interactions are commonly studied with an interest in determining the optimal strategy for prey. However, the implications of deviating from optimal strategy are often unclear. The present study considered these consequences by studying how the direction of an escape response affects the strategy of prey fish. We simulated these interactions with numerical and analytical mathematics and compared our predictions with measurements in zebrafish larvae (Danio rerio), which are preyed upon by adults of the same species. Consistent with existing theory, we treated the minimum distance between predator and prey as the strategic payoff that prey aim to maximize. We found that these interactions may be characterized by three strategic domains that are defined by the speed of predator relative to the prey. The "fast predator" domain occurs when the predator is more than an order of magnitude faster than the prey. The escape direction of the prey had only a small effect on the minimum distance under these conditions. For the "slow predator" domain, when the prey is faster than the predator, we found that differences in direction had no effect on the minimum distance for a broad range of escape angles. This was the regime in which zebrafish were found to operate. In contrast, the optimal escape angle offers a large benefit to the minimum distance in the intermediate strategic domain. Therefore, optimal strategy is most meaningful to prey fish when predators are faster than prey by less than a factor of 10. This demonstrates that the strategy of a prey animal does not matter under certain conditions that are created by the behavior of the predator., (© The Author 2015. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Prey fish possess a remarkable ability to sense and evade an attack from a larger fish. Despite the importance of these events to the biology of fishes, it remains unclear how sensory cues stimulate an effective evasive maneuver. Here, we show that larval zebrafish (Danio rerio) evade predators using an escape response that is stimulated by the water flow generated by an approaching predator. Measurements of the high-speed responses of larvae in the dark to a robotic predator suggest that larvae respond to the subtle flows in front of the predator using the lateral line system. This flow, known as the bow wave, was visualized and modeled with computational fluid dynamics. According to the predictions of the model, larvae direct their escape away from the side of their body exposed to more rapid flow. This suggests that prey fish use a flow reflex that enables predator evasion by generating a directed maneuver at high speed. These findings demonstrate a sensory-motor mechanism that underlies a behavior that is crucial to the ecology and evolution of fishes., (© 2014. Published by The Company of Biologists Ltd.)

The fluid forces that govern propulsion determine the speed and energetic cost of swimming. These hydrodynamics are scale dependent and it is unclear what forces matter to the tremendous diversity of aquatic animals that are between a millimeter and a centimeter in length. Animals at this scale generally operate within the regime of intermediate Reynolds numbers, where both viscous and inertial fluid forces have the potential to play a role in propulsion. The present study aimed to resolve which forces create thrust and drag in the paddling of the water boatman (Corixidae), an animal that spans much of the intermediate regime (10

The characid fish species Astyanax mexicanus offers a classic comparative model for the evolution of sensory systems. Populations of this species evolved in caves and became blind while others remained in streams (i.e. surface fish) and retained a functional visual system. The flow-sensitive lateral line receptors, called superficial neuromasts, are more numerous in cavefish than in surface fish, but it is unclear whether individual neuromasts differ in sensitivity between these populations. The aims of this study were to determine whether the neuromasts in cavefish impart enhanced sensitivity relative to surface fish and to test whether this aids their ability to sense flow in the absence of visual input. Sensitivity was assessed by modeling the mechanics and hydrodynamics of a flow stimulus. This model required that we measure the dimensions of the transparent cupula of a neuromast, which was visualized with fluorescent microspheres. We found that neuromasts within the eye orbit and in the suborbital region were larger and consequently about twice as sensitive in small adult cavefish as in surface fish. Behavioral experiments found that these cavefish, but not surface fish, were attracted to a 35 Hz flow stimulus. These results support the hypothesis that the large superficial neuromasts of small cavefish aid in flow sensing. We conclude that the morphology of the lateral line could have evolved in cavefish to permit foraging in a cave environment.

Insects exchange respiratory gases via an extensive network of tracheal vessels that open to the surface of the body through spiracular valves. Although gas exchange is known to increase with the opening of these spiracles, it is not clear how this event relates to gas flow through the tracheal system. We examined the relationship between respiratory airflow and spiracle activity in a ventilating insect, the hissing cockroach, Gromphadorhina portentosa, to better understand the complexity of insect respiratory function. Using simultaneous video recordings of multiple spiracular valves, we found that abdominal spiracles open and close in unison during periods of ventilation. Additionally, independent recordings of CO2 release from the abdominal and thoracic regions and observations of hyperoxic tracer gas movement indicate that air is drawn into the thoracic spiracles and expelled from the abdominal spiracles. Our video recordings suggest that this unidirectional flow is driven by abdominal contractions that occur when the abdominal spiracles open. The spiracles then close as the abdomen relaxes and fills with air from the thorax. Therefore, the respiratory system of the hissing cockroach functions as a unidirectional pump through the coordinated action of the spiracles and abdominal musculature. This mechanism may be employed by a broad diversity of large insects that respire by active ventilation.

Fish resist being swept downstream by swimming against a current. Mexican blind cavefish (Astyanax fasciatus) exhibit this innate behavior, rheotaxis, without the aid of vision, but it has been debated whether this ability requires sensing flow with the lateral line system. We tested the role of the lateral line by comparing swimming in a flow chamber in a group of cavefish with a compromised lateral line with a control group. Consistent with previous studies, we found that cavefish orient toward flow and more frequently swim upstream at a higher flow speed. We found that these responses to flow were indistinguishable between fish with compromised and functioning lateral line systems. Rheotaxis was also unaltered by exposing fish to varying degrees of turbulence. These results suggest that the sensing of flow is unnecessary for rheotaxis in cavefish. It appears that tactile stimuli provide a sufficient means of executing this behavior in fish and that rheotaxis may not be a major function of the lateral line system.

The ability of fish to evade predators is central to the ecology and evolution of a diversity of species. However, it is largely unclear how prey fish detect predators in order to initiate their escape. We tested whether larval zebrafish (Danio rerio) sense the flow created by adult predators of the same species. When placed together in a cylindrical arena, we found that larvae were able to escape 70% of predator strikes (mean escape probability P(escape)=0.7, N=13). However, when we pharmacologically ablated the flow-sensitive lateral line system, larvae were rarely capable of escape (mean P(escape)=0.05, N=11). In order to explore the rapid events that facilitate a successful escape, we recorded freely swimming predators and prey using a custom-built camera dolly. This device permitted two-dimensional camera motion to manually track prey and record their escape response with high temporal and spatial resolution. These recordings demonstrated that prey were more than 3 times more likely to evade a suction-feeding predator if they responded before (P(escape)=0.53, N=43), rather than after (P(escape)=0.15, N=13), a predator's mouth opened, which is a highly significant difference. Therefore, flow sensing plays an essential role in predator evasion by facilitating a response prior to a predator's strike.

A skeleton amplifies the minute contractions of muscles to animate the body of an animal. The degree that a muscular contraction displaces an appendage is determined by the gearing provided by the joints of a skeleton. Species that move rapidly commonly possess joints with relatively high gears that produce a large output displacement. However, the speed of an appendage can depend on dynamics that obscure how this motion is influenced by the skeleton. The aim of this review is to resolve mechanical principles that govern the relationship between the gearing and speed of skeletal joints. Forward dynamic models of three rapid force-transmission systems were examined with simulations that varied the gearing of a joint. The leg of a locust, the raptorial appendage of a mantis shrimp, and the jaw of a toad are all driven by the conversion of stored elastic energy into kinetic energy. A locust achieves this conversion with high efficiency when it kicks and thereby applies nearly all stored energy into fast movement. This conversion is unaffected by differences in the leverage of the knee joint, as demonstrated by a maximum kicking speed that was found to be independent of gearing. In contrast, the mantis shrimp creates drag as it strikes toward a prey and thereby loses energy. As a consequence, high gears displace the raptorial appendage relatively far and yield slower motion than do low gears. The muscle that opens a toad's jaw also dissipates energy during ballistic capture of prey. This loss of energy is reduced when jaw opening occurs from the slower muscle contraction produced by a high gear within the jaw. Therefore, the speed of these lever systems is dictated by how gearing affects the efficiency of the conversion of potential energy into kinetic energy. In this way, the energetics of force transmission mediate the relationship between the gearing of a skeletal joint and the maximum speed of its motion.

The geometry of an animal's skeleton governs the transmission of force to its appendages. Joints and rigid elements that create a relatively large output displacement per unit input displacement have been considered to be geared for speed, but the relationship between skeletal geometry and speed is largely untested. The present study explored this subject with experiments and mathematical modeling to evaluate how morphological differences in the raptorial appendage of a mantis shrimp (Gonodactylus smithii) affect the speed of its predatory strike. Based on morphological measurements and material testing, we computationally simulated the transmission of the stored elastic energy that powers a strike and the drag that resists this motion. After verifying the model's predictions against measurements of strike impulse, we conducted a series of simulations that varied the linkage geometry, but were provided with a fixed amount of stored elastic energy. We found that a skeletal geometry that creates a large output displacement achieves a slower maximum speed of rotation than a low-displacement system. This is because a large displacement by the appendage causes a relatively large proportion of its elastic energy to be lost to the generation of drag. Therefore, the efficiency of transmission from elastic to kinetic energy mediates the relationship between the geometry and the speed of a skeleton. We propose that transmission efficiency plays a similar role in form-function relationships for skeletal systems in a diversity of animals.

Lever systems within a skeleton transmit force with a capacity determined by the mechanical advantage, A. A is the distance from input force to a joint, divided by the distance from the joint to the output force. A lever with a relatively high A in static equilibrium has a great capacity to generate force but moves a load over a small distance. Therefore, the geometry of a skeletal lever presents a trade-off between force and speed under quasi-static conditions. The present study considers skeletal dynamics that do not assume static equilibrium by modelling kicking by a locust leg, which is powered by stored elastic energy. This model predicts that the output force of this lever is proportional to A, but its maximum speed is independent of A. Therefore, no trade-off between force and velocity exists in a lever system with spring-mass dynamics. This demonstrates that the motion of a skeleton depends on the major forces that govern its dynamics and cannot be inferred from skeletal geometry alone.

Running performance depends on a mechanical interaction between the feet of an animal and the substrate. This interaction may differ between two species of sand lizard from the Mojave Desert that have different locomotor morphologies and habitat distributions. Uma scorparia possesses toe fringes and inhabits dunes, whereas the closely related Callisaurus draconoides lacks fringes and is found on dune and wash habitats. The present study evaluated whether these distribution patterns are related to differential locomotor performance on the fine sand of the dunes and the course sand of the wash habitat. We measured the kinematics of sprinting and characterized differences in grain size distribution and surface strength of the soil in both habitats. Although wash sand had a surface strength (15.4±6.2 kPa) that was more than three times that of dune sand (4.7±2.1 kPa), both species ran with similar sprinting performance on the two types of soil. The broadly distributed C. draconoides ran with a slightly (22%) faster maximum speed (2.2±0.2 m s(-1)) than the dune-dwelling U. scorparia (1.8±0.2 m s(-1)) on dune sand, but not on wash sand. Furthermore, there were no significant differences in maximum acceleration or the time to attain maximum speed between species or between substrates. These results suggest that differences in habitat distribution between these species are not related to locomotor performance and that sprinting ability is dominated neither by environmental differences in substrate nor the presence of toe fringes.

The ability of a predator fish to capture a prey fish depends on the hydrodynamics of the prey and its behavioral response to the predator's strike. Despite the importance of this predator-prey interaction to the ecology and evolution of a diversity of fish, it is unclear what factors dictate a fish's ability to evade capture. The present study evaluated how the specific gravity of a prey fish's body affects the kinematics of prey capture and the signals detected by the lateral line system of the prey during the strike of a suction-feeding predator. The specific gravity of zebrafish (Danio rerio) larvae was measured with high precision from recordings of terminal velocity in solutions of varying density. This novel method found that specific gravity decreased by ∼5% (from 1.063, N=8, to 1.011, N=35) when the swim bladder inflates. To examine the functional consequences of this change, we developed a mathematical model of the hydrodynamics of prey in the flow field created by a suction-feeding predator. This model found that the observed decrease in specific gravity due to swim bladder inflation causes an 80% reduction of the flow velocity around the prey's body. Therefore, swim bladder inflation causes a substantial reduction in the flow signal that may be sensed by the lateral line system to evade capture. These findings demonstrate that the ability of a prey fish to sense a predator depends crucially on the specific gravity of the prey.

Fish use the lateral line system to sense the water flow created by a predator's strike. Despite its potential importance to the survival of a diversity of species, it is unclear whether this ability becomes compromised when a fish swims. Therefore, the present study compared the behavioral responsiveness of swimming and motionless zebrafish (Danio rerio) larvae when exposed to the flow of a suction-feeding predator. This flow was generated with an impulse chamber, which is a device that we developed to generate a repeatable stimulus with a computer-controlled servo motor. Using high-speed video recordings, we found that about three-quarters (0.76, N=121) of motionless larvae responded to the stimulus with an escape response. These larvae were 66% more likely to respond to flow directed perpendicular than flow running parallel to the body. Swimming larvae exhibited a 0.40 response probability and were therefore nearly half as likely to respond to flow as motionless larvae. However, the latency between stimulus and response was unaffected by swimming or the direction of flow. Therefore, swimming creates changes in the hydrodynamics or neurophysiology of a larval fish that diminish the probability, but not the speed, of their response to a flow stimulus. These findings demonstrate a sensory benefit to the intermittent swimming behavior observed among a broad diversity of fishes.

The lateral line system detects water flow, which allows fish to orient their swimming with respect to hydrodynamic cues. However, it is unclear whether this sense plays a role in the control of propulsion. Hydrodynamic theory suggests that fish could reduce drag by coordinating the motion of the head relative to detected flow signals. To test this hypothesis, we performed measurements of undulatory kinematics during steady swimming in the golden shiner (Notemigonus crysoleucas) at three speeds (4.5, 11.0 and 22.0 cm s(-1)). We found that the phase shift between yaw angle and lateral velocity (20.5+/-13.1 deg., N=5) was significantly greater than the theoretical optimum (0 deg.) and the amplitude of these variables created a hydrodynamic index (H=0.05+/-0.03, N=6) that was less than an order of magnitude below the theoretical prediction. Furthermore, we repeated these measurements after pharmacologically ablating the lateral line hair cells and found that drag reduction was not adversely influenced by disabling the lateral line system. Therefore, flow sensing does not facilitate active drag reduction. However, we discovered that ablating the lateral line causes the envelope of lateral displacement to nearly double at the envelope's most narrow point for swimming at 4.5 cm s(-1). Therefore, fish may use hydrodynamic sensing to modulate the lateral amplitude of slow undulatory swimming, which could allow rapid responses to changes in environmental flow.

Hair cells of the lateral line system in fish may differ in their susceptibility to damage by aminoglycoside antibiotics. Gentamicin has been reported to damage hair cells within canal neuromasts, but not those within superficial neuromasts. This finding, based on SEM imaging, indicates a distinction in the physiology of hair cells between the two classes of neuromast. Studies concerned with the individual roles of canal and superficial neuromasts in behavior have taken advantage of this effect in an attempt to selectively disable canal neuromasts without affecting superficial neuromast function. Here we present an experimental test of the hypothesis that canal neuromasts are more vulnerable to gentamicin than superficial neuromasts. We measured the effect of gentamicin exposure on hair cells using vital stains (DASPEI and FM1-43) in the neuromasts of Mexican blind cave fish (Astyanaxfasciatus) and zebrafish (Daniorerio). Contrary to the findings of prior studies that used SEM, gentamicin significantly reduced dye uptake by hair cells of both canal and superficial neuromasts in both species. Therefore, lateral line hair cells of both neuromast types are vulnerable to gentamicin ototoxicity. These findings argue for a re-evaluation of the results of studies that have used gentamicin to differentiate the roles of the two classes of neuromast in fish behavior., (Published by Elsevier B.V.)

Fish exhibit many behaviors that involve sensing water flows with their lateral-line system. In many situations, viscosity affects how the flow interacts with the body of the fish and the neuromasts of the lateral line. Here we discuss how viscosity influences the stimulus to the fish lateral-line system. The movement of a fish's body creates flows that can interfere with the detection of external signals, but these flows can also serve as a source of information about nearby obstacles and the fish's own hydrodynamic performance. The viscous boundary layer on the surface of the skin alters external signals by attenuating the low-frequency components of stimuli. The stimulus to each neuromast depends on the interaction of the fluid surrounding the neuromast and the structural properties of that neuromast, including the number of mechanosensory hair cells it contains. A consideration of the influences of viscosity on flow, at both the whole-body and receptor levels, offers the promise of a more comprehensive understanding of the signals involved in behaviors mediated by the lateral-line system.

Larval fishes have a remarkable ability to sense and evade the feeding strike of a predator fish with a rapid escape manoeuvre. Although the neuromuscular control of this behaviour is well studied, it is not clear what stimulus allows a larva to sense a predator. Here we show that this escape response is triggered by the water flow created during a predator's strike. Using a novel device, the impulse chamber, zebrafish (Danio rerio) larvae were exposed to this accelerating flow with high repeatability. Larvae responded to this stimulus with an escape response having a latency (mode=13-15 ms) that was fast enough to respond to predators. This flow was detected by the lateral line system, which includes mechanosensory hair cells within the skin. Pharmacologically ablating these cells caused the escape response to diminish, but then recover as the hair cells regenerated. These findings demonstrate that the lateral line system plays a role in predator evasion at this vulnerable stage of growth in fishes.

A great diversity of aquatic animals detects water flow with ciliated mechanoreceptors on the body's surface. In order to understand how these receptors mechanically filter signals, we developed a theoretical model of the superficial neuromast in the fish lateral line system. The cupula of the neuromast was modeled as a cylindrical beam that deflects in response to an oscillating flow field. Its accuracy was verified by comparison with prior measurements of cupular deflection in larval zebrafish (Danio rerio). The model predicts that the boundary layer of flow over the body attenuates low-frequency stimuli. The fluid-structure interaction between this flow and the cupula attenuates high-frequency stimuli. The number and height of hair cell kinocilia and the dimensions of the cupular matrix determine the range of intermediate frequencies to which a neuromast is sensitive. By articulating the individual mechanical contributions of the boundary layer and the components of cupular morphology, this model provides the theoretical framework for understanding how a hydrodynamic receptor filters flow signals.

The lateral line system of fish and amphibians detects water flow with receptors on the surface of the body. Although differences in the shape of these receptors, called neuromasts, are known to influence their mechanics, it is unclear how neuromast morphology affects the sensitivity of the lateral line system. We examined the functional consequences of morphological variation by measuring the dimensions of superficial neuromasts in zebrafish larvae (Danio rerio) and mathematically modeling their mechanics. These measurements used a novel morphometric technique that recorded landmarks in three dimensions at a microscopic scale. The mathematical model predicted mechanical sensitivity as the ratio of neuromast deflection to flow velocity for a range of stimulus frequencies. These predictions suggest that variation in morphology within this species generates a greater than 30-fold range in the amplitude of sensitivity and more than a 200-fold range of variation in cut-off frequency. Most of this variation was generated by differences in neuromast height that do not correlate with body position. Our results suggest that natural variation in cupular height within a species is capable of generating large differences in their mechanical filtering and dynamic range.

Superficial neuromasts are structures that detect water flow on the surface of the body of fish and amphibians. As a component of the lateral line system, these receptors are distributed along the body, where they sense flow patterns that mediate a wide variety of behaviors. Their ability to detect flow is governed by their structural properties, yet the micromechanics of superficial neuromasts are not well understood. The aim of this study was to examine these mechanics in zebrafish (Danio rerio) larvae by measuring the flexural stiffness of individual neuromasts. Each neuromast possesses a gelatinous cupula that is anchored to hair cells by kinocilia. Using quasi-static bending tests of the proximal region of the cupula, we found that flexural stiffness is proportional to the number of hair cells, and consequently the number of kinocilia, within a neuromast. From this relationship, the flexural stiffness of an individual kinocilium was found to be 2.4 x 10(-20) N m2. Using this value, we estimate that the 11 kinocilia in an average cupula generate more than four-fifths of the total flexural stiffness in the proximal region. The relatively minor contribution of the cupular matrix may be attributed to its highly compliant material composition (Young's modulus of approximately 21 Pa). The distal tip of the cupula is entirely composed of this material and is consequently predicted to be at least an order of magnitude more flexible than the proximal region. These findings suggest that the transduction of flow by a superficial neuromast depends on structural dynamics that are dominated by the number and height of kinocilia.

Studying the mechanics of swirling water has solved a mystery about the evolution of body shape and size in jellyfish.

Many fish species transform in body shape during growth, but it remains unclear how this influences the mechanics of locomotion. Therefore, the present study focused on understanding how drag generation during coasting is affected by ontogenetic changes in the morphology of zebrafish (Danio rerio). The shapes of the body and fins were measured from photographs of fish ranging in size from small larvae to mature adults and these morphometrics were compared to drag coefficients calculated from high-speed video recordings of routine swimming. We found that the viscous drag coefficient of larval and juvenile fish increased by more than an order of magnitude during growth and the inertial drag coefficient decreased at a comparable rate in adults. These hydrodynamic changes occurred as zebrafish disproportionately increased the span of their fins and their body changed shape from elongated to streamlined, as reflected by the logistic growth of a newly defined streamlining index, SL. These results suggest that morphological changes incur a performance cost by generating greater drag when larvae and juveniles operate in the viscous regime, but later provide a performance benefit by reducing pressure drag in the inertial regime of the adult stage., ((c) 2006 Wiley-Liss, Inc.)