skip to main content
US FlagAn official website of the United States government
dot gov icon
Official websites use .gov
A .gov website belongs to an official government organization in the United States.
https lock icon
Secure .gov websites use HTTPS
A lock ( lock ) or https:// means you've safely connected to the .gov website. Share sensitive information only on official, secure websites.


Title: 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.  more » « less
Award ID(s):
1856237 1907156 2102891
PAR ID:
10337194
Author(s) / Creator(s):
; ; ; ; ; ;
Date Published:
Journal Name:
Proceedings of the National Academy of Sciences
Volume:
118
Issue:
49
ISSN:
0027-8424
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract Many aquatic animals swim by undulatory body movements and understanding the diversity of these movements could unlock the potential for designing better underwater robots. Here, we analyzed the steady swimming kinematics of a diverse group of fish species to investigate whether their undulatory movements can be represented using a series of interconnected multi-segment models, and if so, to identify the key factors driving the segment configuration of the models. Our results show that the steady swimming kinematics of fishes can be described successfully using parsimonious models, 83% of which had fewer than five segments. In these models, the anterior segments were significantly longer than the posterior segments, and there was a direct link between segment configuration and swimming kinematics, body shape, and Reynolds number. The models representing eel-like fishes with elongated bodies and fishes swimming at high Reynolds numbers had more segments and less segment length variability along the body than the models representing other fishes. These fishes recruited their anterior bodies to a greater extent, initiating the undulatory wave more anteriorly. Two shape parameters, related to axial and overall body thickness, predicted segment configuration with moderate to high success rate. We found that head morphology was a good predictor of its segment length. While there was a large variation in head segments, the length of tail segments was similar across all models. Given that fishes exhibited variable caudal fin shapes, the consistency of tail segments could be a result of an evolutionary constraint tuned for high propulsive efficiency. The bio-inspired multi-segment models presented in this study highlight the key bending points along the body and can be used to decide on the placement of actuators in fish-inspired robots, to model hydrodynamic forces in theoretical and computational studies, or for predicting muscle activation patterns during swimming. 
    more » « less
  2. Abstract The epaulette shark, Hemiscyllium ocellatum, is a small, reef-dwelling, benthic shark that—using its paired fins—can walk, both in and out of water. Within the reef flats, this species experiences short periods of elevated CO2 and hypoxia as well as fluctuating temperatures as reef flats become isolated with the outgoing tide. Past studies have shown that this species is robust (i.e., respiratory and metabolic performance, behavior) to climate change-relevant elevated CO2 levels as well as hypoxia and anoxia tolerant. However, epaulette shark embryos reared under ocean warming conditions hatch earlier and smaller, with altered patterns and coloration, and with higher metabolic costs than their current-day counterparts. Findings to date suggest that this species has adaptations to tolerate some, but perhaps not all, of the challenging conditions predicted for the 21st century. As such, the epaulette shark is emerging as a model system to understand vertebrate physiology in changing oceans. Yet, few studies have investigated the kinematics of walking and swimming, which may be vital to their biological fitness, considering their habitat and propensity for challenging environmental conditions. Given that neonates retain embryonic nutrition via an internalized yolk sac, resulting in a bulbous abdomen, while juveniles actively forage for worms, crustaceans, and small fishes, we hypothesized that difference in body shape over early ontogeny would affect locomotor performance. To test this, we examined neonate and juvenile locomotor kinematics during the three aquatic gaits they utilize—slow-to-medium walking, fast walking, and swimming—using 13 anatomical landmarks along the fins, girdles, and body midline. We found that differences in body shape did not alter kinematics between neonates and juveniles. Overall velocity, fin rotation, axial bending, and tail beat frequency and amplitude were consistent between early life stages. Data suggest that the locomotor kinematics are maintained between neonate and juvenile epaulette sharks, even as their feeding strategy changes. Studying epaulette shark locomotion allows us to understand this—and perhaps related—species’ ability to move within and away from challenging conditions in their habitats. Such locomotor traits may not only be key to survival, in general, as a small, benthic mesopredator (i.e., movements required to maneuver into small reef crevices to avoid aerial and aquatic predators), but also be related to their sustained physiological performance under challenging environmental conditions, including those associated with climate change—a topic worthy of future investigation. 
    more » « less
  3. High efficiency lunate-tail swimming with high-aspect-ratio lifting surfaces has evolved in many vertebrate lineages, from fish to cetaceans. Baleen whales (Mysticeti) are the largest swimming animals that exhibit this locomotor strategy, and present an ideal study system to examine how morphology and the kinematics of swimming scale to the largest body sizes. We used data from whale-borne inertial sensors coupled with morphometric measurements from aerial drones to calculate the hydrodynamic performance of oscillatory swimming in six baleen whale species ranging in body length from 5 to 25 m (fin whale, Balaenoptera physalus; Bryde’s whale, Balaenoptera edeni; sei whale, Balaenoptera borealis; Antarctic minke whale, Balaenoptera bonaerensis; humpback whale, Megaptera novaeangliae; and blue whale, Balaenoptera musculus). We found that mass-specific thrust increased with both swimming speed and body size. Froude efficiency, defined as the ratio of useful power output to the rate of energy input (Sloop, 1978), generally increased with swimming speed but decreased on average with increasing body size. This finding is contrary to previous results in smaller animals, where Froude efficiency increased with body size. Although our empirically parameterized estimates for swimming baleen whale drag were higher than those of a simple gliding model, oscillatory locomotion at this scale exhibits generally high Froude efficiency as in other adept swimmers. Our results quantify the fine- scale kinematics and estimate the hydrodynamics of routine and energetically expensive swimming modes at the largest scale. 
    more » « less
  4. null (Ed.)
    Biological armours are potent model systems for understanding the complex series of competing demands on protective exoskeletons; after all, armoured organisms are the product of millions of years of refined engineering under the harshest conditions. Fishes are no strangers to armour, with various types of armour plating common to the 400–500 Myr of evolution in both jawed and jawless fishes. Here, we focus on the poachers (Agonidae), a family of armoured fishes native to temperate waters of the Pacific rim. We examined armour morphology, body stiffness and swimming performance in the northern spearnose poacher ( Agonopsis vulsa ) over ontogeny. As juveniles, these fishes make frequent nocturnal forays into the water column in search of food, while heavily armoured adults are bound to the benthos. Most armour dimensions and density increase with body length, as does body stiffness. Juvenile poachers have enlarged spines on their armour whereas adults invest more mineral in armour plate bases. Adults are stiffer and accelerate faster than juveniles with an anguilliform swimming mode. Subadults more closely approximate adults more than smaller juveniles, with regards to both swimming and armour mechanics. Poacher armour serves multiple functions over ontogeny, from facilitating locomotion, slowing sinking and providing defence. 
    more » « less
  5. 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. 
    more » « less