Convergence of undulatory swimming kinematics across a diversity of fishes

Fishes exhibit an astounding diversity of locomotor behaviors from classic swimming with their body and fins to jumping, flying, walking, and burrowing. Fishes that use their body and caudal fin (BCF) during undulatory swimming have been traditionally divided into modes based on the length of the propulsive body wave and the ratio of head:tail oscillation amplitude: anguilliform, subcarangiform, carangiform, and thunniform. This classification was first proposed based on key morphological traits, such as body stiffness and elongation, to group fishes based on their expected swimming mechanics. Here, we present a comparative study of 44 diverse species quantifying the kinematics and morphology of BCF-swimming fishes. Our results reveal that most species we studied share similar oscillation amplitude during steady locomotion that can be modeled using a second-degree order polynomial. The length of the propulsive body wave was shorter for species classified as anguilliform and longer for those classified as thunniform, although substantial variability existed both within and among species. Moreover, there was no decrease in head:tail amplitude from the anguilliform to thunniform mode of locomotion as we expected from the traditional classification. While the expected swimming modes correlated with morphological traits, they did not accurately represent the kinematics of BCF locomotion. These results indicate that even fish species differing as substantially in morphology as tuna and eel exhibit statistically similar two-dimensional midline kinematics and point toward unifying locomotor hydrodynamic mechanisms that can serve as the basis for understanding aquatic locomotion and controlling biomimetic aquatic robots.

Kinematic Analysis During Straight Line Free Swimming in Horses: Part 1 - Forelimbs

Background: Swimming is used for rehabilitation and conditioning purposes in equine sports medicine despite the lack of understanding of equine swimming kinematics. The aim of this study was to assess forelimb joints kinematics (elbow, carpus, and fetlock) in swimming horses. The specific objectives were 1- to calculate and compare joint angles in swimming vs. passive mobilizations (PM), 2- to determine joint angular velocities during a swimming stride cycle.Methods: Eleven elite endurance horses swam in a 100-m straight pool. Underwater (swimming) and overground (PM) videos were recorded from the horses' left side. Joint markers were applied on the lateral hoof wall, lateral metacarpal epicondyle, ulnar carpal bone, lateral humeral epicondyle, and the greater tubercle of humerus, from which elbow, carpus and fetlock angles, and angular velocities were obtained. As a reference, maximal fetlock, carpus, and elbow flexion/extension angles were determined during PM overground. Differences between angle extrema, angular velocities and range of motion (ROM) were compared.Results: Carpus and fetlock ROM were significantly smaller (p < 0.001) during swimming when compared with PM, while there was no difference in elbow ROM between both situations. The carpus had the greatest ROM of all joints during swimming. Absolute angular velocities values of all joints during swimming were greater during retraction than protraction (p < 0.001). When compared to other joints during protraction, the carpus joint reached the highest angular velocity.Conclusion: Swimming, as a rehabilitation exercise, has the potential to benefit horses where great elbow ROM with a moderate carpus and fetlock extension are wanted.

A comparison of stride parameters and carpal and tarsal joint angles during terrestrial and swimming locomotion in domestic dogs

In recent years, canine hydrotherapy has become increasingly popular to treat a range of conditions despite a lack of empirical evidence. It is currently unclear whether joint angles and limb movements performed by dogs during swimming are quantifiably beneficial for healthy animals. This study investigated the swimming kinematics of healthy dogs to establish baseline data for this activity and compare limb kinematics to that of overground locomotion. Kinematic data were recorded from eight healthy dolichocephalic dogs (mean age: 3.4±2.2) of a variety of breeds. Overground data were collected prior to swimming and consisted of dogs trotting on a flat surface. Swimming data were collected using an underwater camera during a standard hydrotherapy session conducted by a trained canine hydrotherapist. Range of motion, primarily due to an increase in flexion, was significantly greater (P<0.005) during swimming than trotting. Stride length (P<0.001) and frequency (P<0.005) were both significantly reduced in swimming compared to trot. Swimming kinematics recorded in this study are consistent with previously published data on canine aquatic locomotion but differ from those previously reported for water treadmill exercise. This study provides an insight into aquatic locomotion in healthy dogs indicating that range of motion exceeds that of terrestrial gaits. It is unclear whether these changes are beneficial for healthy animals and therefore further research is required to develop evidence-based protocols for industry practice.

Swimming kinematics of cyprids of the barnacle Balanus glandula

Larvae of barnacles typically pass through naupliar and cyprid planktonic stages before settlement and metamorphosis. As the final larval stage, cyprids swim much faster than nauplii and in turbulent fluid environments with high shears as they seek habitat. Cyprids swim with six pairs of reciprocating thoracic appendages and use two anterior antennules during settlement. Our understanding of how thoracic appendages generate movement is limited due to short stroke intervals (∼5 ms) that impede observations of the shape and trajectory of appendages. Here, we used high speed videography to observe both free-swimming and tethered cyprids of the intertidal acorn barnacle Balanus glandula to produce a comprehensive description of thoracic appendage swimming kinematics. Cyprids used a drag-based method of swimming: their six pairs of thoracic appendages moved through metachronal power strokes and synchronous recovery strokes similar to the thoracopod motions in calanoid copepods during escape swimming. During the power stroke, plumose setae on each appendage pair spread laterally into a high surface area and high drag paddle composed of a meshwork of fused setules. This interconnected setal array collapsed into a low surface area and low drag shape during the recovery stroke. These effective swimming appendages allowed cyprids to move upwards at an average speed of 1.4 cm s−1 (about 25 body lengths s−1) with an average beat frequency of 16 beats s−1, and reach an instantaneous velocity of up to 6 cm s−1. Beat frequency of the thoracic appendages was significantly associated with speed, with higher beat frequencies indicating faster swimming speed. At their average speed, cyprids moved at the intermediate Reynolds number of ∼10, in which both viscous and inertial forces affected movement. Cyprids could alter swimming direction by sweeping the posterior-most appendage pair to one side and beating the remaining thoracic appendages synchronously through the power stroke with greater motion on the outside of their turn. These results greatly enhance our understanding both of cyprid motility and how small planktonic organisms can use swimming appendages with fused setule arrays to reach high swimming speeds and affect directional changes.

Swimming kinematics and performance of spinal transected lampreys with different levels of axon regeneration

Neural and functional recovery in lampreys from spinal cord transection has been well documented. However, the extent of axon regeneration is highly variable and it is not known whether it is related to the level of behavioral recovery. To address this, we examined how swimming kinematics were related to axon regeneration by quantifying the relationship between swimming performance and percent axon regeneration of transected lampreys after 11 weeks of recovery. We found that swimming speed was not related to percent axon regeneration but it was closely related to body wave frequency and speed. However, wave frequency and speed varied greatly within individuals which resulted in swimming speed also varying within individuals. In fact, most recovered individuals, regardless of percent axon regeneration, could swim at fast and slow speeds. However, none of the transected individuals were able to generate body waves as large as the control lampreys. In order to swim faster, transected lampreys increased their wave frequencies and, as a result, transected lampreys had much higher frequencies than control lamprey at comparable swimming velocities. These data suggest that the control lampreys swam more efficiently than transected lampreys. In conclusion, there appears to be a minimal recovery threshold in terms of percent axon regeneration required for lampreys to be capable of swimming, however, there also seems to be a limit to how much they can behaviorally recover.

The response of the copepod Acartia tonsa to the hydrodynamic cues of small-scale, dissipative eddies in turbulence

This study quantifies the behavioral response of a marine copepod (Acartia tonsa) to individual, small-scale, dissipative vortices that are ubiquitous in turbulence. Vortex structures were created in the laboratory using a physical model of a Burgers vortex with characteristics corresponding to typical dissipative vortices that copepods are likely to encounter in the turbulent cascade. To examine the directional response of copepods, vortices were generated with the vortex axis aligned in either horizontal or vertical directions. Tomographic particle image velocimetry was used to measure the volumetric velocity field of the vortex. Three-dimensional copepod trajectories were digitally reconstructed and overlaid on the vortex flow field to quantify A. tonsa’s swimming kinematics relative to the velocity field and to provide insight to the copepod behavioral response to hydrodynamic cues. The data show significant changes in swimming kinematics and an increase in relative swimming velocity and hop frequency with increasing vortex strength. Furthermore, in moderate-to-strong vortices, A. tonsa moved at elevated speed in the same direction as the swirling flow and followed spiral trajectories around the vortex, which would retain the copepod within the feature and increase encounter rates with other similarly behaving Acartia. While changes in swimming kinematics depended on vortex intensity, orientation of the vortex axis showed minimal significant effect. Hop and escape jump densities were largest in the vortex core, which is spatially coincident with the peak in vorticity suggesting that vorticity is the hydrodynamic cue that evokes these behaviors.

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

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.

Swimming kinematics and hydrodynamics of barnacle larvae throughout development

Changes in size strongly influence organisms' ecological performances. For aquatic organisms, they can transition from viscosity- to inertia-dominated fluid regimes as they grow. Such transitions are often associated with changes in morphology, swimming speed and kinematics. Barnacles do not fit into this norm as they have two morphologically distinct planktonic larval phases that swim differently but are of comparable sizes and operate in the same fluid regime (Reynolds number 10 0 –10 1 ). We quantified the hydrodynamics of the rocky intertidal stalked barnacle Capitulum mitella from the nauplius II to cyprid stage and examined how kinematics and size increases affect its swimming performance. Cyprids beat their appendages in a metachronal wave to swim faster, more smoothly, and with less backwards slip per beat cycle than did all naupliar stages. Micro-particle image velocimetry showed that cyprids generated trailing viscous vortex rings that pushed water backwards for propulsion, contrary to the nauplii's forward suction current for particle capture. Our observations highlight that zooplankton swimming performance can shift via morphological and kinematic modifications without a significant size increase. The divergence in ecological functions through ontogeny in barnacles and the removal of feeding requirement likely contributed to the evolution of the specialized, taxonomically unique cyprid phase.