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1. Volitional Swimming Kinematics of Blacktip Sharks, Carcharhinus limbatus, in the Wild

   https://doi.org/10.3390/drones4040078  

   Porter, Marianne E.; Ruddy, Braden T.; Kajiura, Stephen M. (December 2020, Drones)

   null (Ed.)

   Recent work showed that two species of hammerhead sharks operated as a double oscillating system, where frequency and amplitude differed in the anterior and posterior parts of the body. We hypothesized that a double oscillating system would be present in a large, volitionally swimming, conventionally shaped carcharhinid shark. Swimming kinematics analyses provide quantification to mechanistically examine swimming within and among species. Here, we quantify blacktip shark (Carcharhinus limbatus) volitional swimming kinematics under natural conditions to assess variation between anterior and posterior body regions and demonstrate the presence of a double oscillating system. We captured footage of 80 individual blacktips swimming in the wild using a DJI Phantom 4 Pro aerial drone. The widespread accessibility of aerial drone technology has allowed for greater observation of wild marine megafauna. We used Loggerpro motion tracking software to track five anatomical landmarks frame by frame to calculate tailbeat frequency, tailbeat amplitude, speed, and anterior/posterior variables: amplitude and frequency of the head and tail, and the body curvature measured as anterior and posterior flexion. We found significant increases in tailbeat frequency and amplitude with increasing swimming speed. Tailbeat frequency decreased and tailbeat amplitude increased as posterior flexion amplitude increased. We found significant differences between anterior and posterior amplitudes and frequencies, suggesting a double oscillating modality of wave propagation. These data support previous work that hypothesized the importance of a double oscillating system for increased sensory perception. These methods demonstrate the utility of quantifying swimming kinematics of wild animals through direct observation, with the potential to apply a biomechanical perspective to movement ecology paradigms. 

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2. Regulation of the swimming kinematics of lampreys Petromyzon marinus across changes in viscosity

   https://doi.org/10.1242/jeb.245457  

   Tytell, Eric D.; Cooper, Lauren O.; Lin, Yuexia Luna; Reis, Pedro M. (May 2023, Journal of Experimental Biology)

   ABSTRACT The bodies of most swimming fishes are very flexible and deform as result of both external fluid dynamic forces and internal musculoskeletal forces. If fluid forces change, the body motion will also change unless the fish senses the change and alters its muscle activity to compensate. Lampreys and other fishes have mechanosensory cells in their spinal cords that allow them to sense how their body is bending. We hypothesized that lampreys (Petromyzon marinus) actively regulate body curvature to maintain a fairly constant swimming waveform even as swimming speed and fluid dynamic forces change. To test this hypothesis, we measured the steady swimming kinematics of lampreys swimming in normal water, and water in which the viscosity was increased by 10 or 20 times by adding methylcellulose. Increasing the viscosity over this range increases the drag coefficient, potentially increasing fluid forces up to 40%. Previous computational results suggested that if lampreys did not compensate for these forces, the swimming speed would drop by about 52%, the amplitude would drop by 39%, and posterior body curvature would increase by about 31%, while tail beat frequency would remain the same. Five juvenile sea lampreys were filmed swimming through still water, and midlines were digitized using standard techniques. Although swimming speed dropped by 44% from 1× to 10× viscosity, amplitude only decreased by 4%, and curvature increased by 7%, a much smaller change than the amount we estimated if there was no compensation. To examine the waveform overall, we performed a complex orthogonal decomposition and found that the first mode of the swimming waveform (the primary swimming pattern) did not change substantially, even at 20× viscosity. Thus, it appears that lampreys are compensating, at least partially, for the changes in viscosity, which in turn suggests that sensory feedback is involved in regulating the body waveform. 

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3. Intermittent propulsion in largemouth bass, Micropterus salmoides , increases power production at low swimming speeds

   https://doi.org/10.1098/rsbl.2021.0658  

   Coughlin, D. J.; Chrostek, J. D.; Ellerby, D. J. (May 2022, Biology Letters)

   Locomotion dominates animal energy budgets, and selection should favour behaviours that minimize transportation costs. Recent fieldwork has altered our understanding of the preferred modes of locomotion in fishes. For instance, bluegill employ a sustainable intermittent swimming form with 2–3 tail beats alternating with short glides. Volitional swimming studies in the laboratory with bluegill suggest that the propulsive phase reflects a fixed-gear constraint on body–caudal-fin activity. Largemouth bass ( Micropterus salmoides ) also reportedly display intermittent swimming in the field. We examined swimming by bass in a static tank to quantify the parameters of volitional locomotion, including tailbeat frequency and glide duration, across a range of swimming speeds. We found that tailbeat frequency was not related to speed at low swimming speeds. Instead, speed was a function of glide duration between propulsive events, with glide duration decreasing as speed increased. The propulsive Strouhal number remained within the range that maximizes propulsive efficiency. We used muscle mechanics experiments to simulate power production by muscle operating under intermittent versus steady conditions. Workloop data suggest that intermittent activity allows fish to swim efficiently and avoid the drag-induced greater energetic cost of continuous swimming. The results offer support for a new perspective on fish locomotion: intermittent swimming is crucial to aerobic swimming energetics. 

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4. A Tail of Four Fishes: An analysis of kinematics and material properties of elongate fishes

   https://doi.org/10.1093/icb/icab060  

   Naughton, Lydia F; Kruppert, Sebastian; Jackson, Beverly; Porter, Marianne E; Donatelli, Cassandra M (May 2021, Integrative and Comparative Biology)

   null (Ed.)

   Abstract The elongate body plan is present in many groups of fishes, and this morphology dictates functional consequences seen in swimming behavior. Previous work has shown that increasing the number of vertebrae, or decreasing the intervertebral joint length, in a fixed length artificial system increases stiffness. Tails with increased stiffness can generate more power from tail beats, resulting in an increased mean swimming speed. This demonstrates the impacts of morphology on both material properties and kinematics, establishing mechanisms for form contributing to function. Here, we wanted to investigate relationships between form and ecological function, such as differences in dietary strategies and habitat preferences among fish species. This study aims to characterize and compare the kinematics, material properties, and vertebral morphology of four species of elongate fishes: Anoplarchus insignis, Anoplarchus purpurescens, Xiphister atropurpureus, and Xiphister mucosus. We hypothesized that these properties would differ among the four species due to their differential ecological niches. To calculate kinematic variables, we filmed these fishes swimming volitionally. We also measured body stiffness by bending the abdominal and tail regions of sacrificed individuals in different stages of dissection (whole body, removed skin, removed muscle). Finally, we counted the number of vertebrae from CT scans of each species to quantify vertebral morphology. Principal component and linear discriminant analyses suggested that the elongate fish species can be distinguished from one another by their material properties, morphology, and swimming kinematics. With this information combined, we can draw connections between the physical properties of the fishes and their ecological niches. 

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5. Fish-inspired segment models for undulatory steady swimming

   https://doi.org/10.1088/1748-3190/ac6bd6  

   Akanyeti, Otar; Di Santo, Valentina; Goerig, Elsa; Wainwright, Dylan K; Liao, James C; Castro-Santos, Theodore; Lauder, George V (May 2022, Bioinspiration & Biomimetics)

   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. 

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