Swimming Mode Research Articles

Towards passive station holding of autonomous underwater vehicles inspired by fish behaviour in unsteady flows

Some species of fish are able to alter their mode of swimming to interact with naturally produced vortices; the use of these gaits reduces the energy expended by the fish. To analyse the feasibility of autonomous underwater vehicles replicating these gaits, a series of experiments are performed with unpowered rigid and flexible bodies positioned in the Kármán wake of a rigid cylinder. Simple motion capture techniques are used to capture the bodies’ lateral and upstream motion in the flow. The results demonstrate that manufactured bodies are capable of passively mimicking fish behaviours, to a limited extent. More importantly, it was concluded that while significant upstream movement was possible for a manufactured object, it was achievable irrespective of the stiffness of the material. For autonomous underwater vehicles operating in unsteady flow regimes, an ability to utilise energy-saving gaits may improve the range or operational time.

Implementations of the route planning scenarios for the autonomous robotic fish with the optimized propulsion mechanism

Abstract Various human problems are tried to resolve with biomimetic design which imitate biological forms. A biomimetic Carangiform robotic fish provides great benefits with flexible maneuverability, high propulsion efficiency and less noisy considering classical rotary underwater vehicles. This paper presents a dynamic simulation model of the Carangiform robotic fish with flexible multi-joint propulsion mechanism considered as an artificial spine system for two swimming cases. In order to swim like a real fish, multi-joint propulsion mechanism assumed a series planar hinge joints which represent vertebras is adjusted by optimizing with a new searching method which provides precise values as direct search methods. The flapping frequency and the speed are proportional with the tail link lengths and angles of the joints. Thus, the optimization parameters are selected as end point coordinates of the joints and lengths of the each link to imitate the real traveling body wave. Two possible route planning scenarios for the robotic fish model inspired from the Carangiform motion are performed. These scenarios are summarized by two cases. Case 1 is the free swimming mode permits to go straight forward until it faces an obstacle. The fish decides to the turning direction by using decision-making process when it encounters an obstacle and finds the way to turn. In the Case 2, the fish proposes to reach the destination area along the shortest path. When faced with obstacles, it overcomes obstacles and tries to reach the target in the shortest way again.

Reynolds-number scaling of vortex pinch-off on low-aspect-ratio propulsors

Impulsively started, low-aspect-ratio elliptical flat plates have been investigated experimentally to understand the vortex pinch-off dynamics at transitional and fully turbulent Reynolds numbers. The range of Reynolds numbers investigated is representative of those observed in animals that employ rowing and paddling modes of drag-based propulsion and manoeuvring. Elliptical flat plates with five aspect ratios ranging from one to two have been considered, as abstractions of propulsor planforms found in nature. It has been shown that Reynolds-number scaling is primarily determined by plate aspect ratio in terms of both drag forces and vortex pinch-off. Due to vortex-ring growth time scales that are longer than those associated with the development of flow instabilities, the scaling of drag is Reynolds-number-dependent for the aspect-ratio-one flat plate. With increasing aspect ratio, the Reynolds-number dependency decreases as a result of the shorter growth time scales associated with high-aspect-ratio elliptical vortex rings. Large drag peaks are observed during early-stage vortex growth for the higher-aspect-ratio flat plates. The collapse of these peaks with Reynolds number provides insight into the evolutionary convergence process of propulsor planforms used in drag-based swimming modes over diverse scales towards aspect ratios greater than one.

Similarities and Differences for Swimming in Larval and Adult Lampreys.

The spinal locomotor networks controlling swimming behavior in larval and adult lampreys may have some important differences. As an initial step in comparing the locomotor systems in lampreys, in larval animals the relative timing of locomotor movements and muscle burst activity were determined and compared to those previously published for adults. In addition, the kinematics for free swimming in larval and adult lampreys was compared in detail for the first time. First, for swimming in larval animals, the neuromechanical phase lag between the onsets or terminations of muscle burst activity and maximum concave curvature of the body increased with increasing distance along the body, similar to that previously shown in adults. Second, in larval lampreys, but not adults, absolute swimming speed (U; mm s(-1)) increased with animal length (L). In contrast, normalized swimming speed (U'; body lengths [bl] s(-1)) did not increase with L in larval or adult animals. In both larval and adult lampreys, U' and normalized wave speed (V') increased with increasing tail-beat frequency. Wavelength and mechanical phase lag did not vary significantly with tail-beat frequency but were significantly different in larval and adult animals. Swimming in larval animals was characterized by a smaller U/V ratio, Froude efficiency, and Strouhal number than in adults, suggesting less efficient swimming for larval animals. In addition, during swimming in larval lampreys, normalized lateral head movements were larger and normalized lateral tail movements were smaller than for adults. Finally, larval animals had proportionally smaller lateral surface areas of the caudal body and fin areas than adults. These differences are well suited for larval sea lampreys that spend most of the time buried in mud/sand, in which swimming efficiency is not critical, compared to adults that would experience significant selection pressure to evolve higher-efficiency swimming to catch up to and attach to fish for feeding as well as engage in long-distance migration during spawning. Finally, the differences in swim efficiency for larval and adult lampreys are compared to other animals employing the anguilliform mode of swimming.

Diversity of Lipid Distribution in Fish Skeletal Muscle.

Adipose tissue is a lipid storage organ characterized by the pronounced accumulation of adipocytes. Although adipose tissues are found in various parts of the vertebrate body, it is unclear whether these tissues have a common ancestral origin or have evolved in several phylogenetic lineages by independent adipocyte accumulation events. To gain insight into the evolutionary history of vertebrate adipose tissues, we determined the distribution of adipocytes by oil red O staining in skeletal muscle of 10 teleost species spanning eight orders: Tetraodontiformes, Pleuronectiformes, Spariformes, Salmoniformes, Clupeiformes, Beloniformes, Osmeriformes, and Cypriniformes. Accumulation of adipocytes in the myoseptum was observed in many species, including red seabream, rainbow trout, Pacific herring, Pacific saury, zebrafish and giant danio. We also found some order-, species-, and swimming mode-specific distribution patterns of adipocytes: 1) almost complete absence of intramuscular adipocytes in the order Tetraodontiformes (torafugu and spotted green pufferfish), 2) clear adipocyte accumulation in the inclinator muscles of fin in Japanese flounder, 3) a large intramuscular adipose tissue at the root of the dorsal fin in ayu, and 4) thick lipid layers consisting of subcutaneous adipose tissue and red muscle lipids in pelagic migratory fish (Pacific herring and Pacific saury). Of note, Pacific herring and Pacific saury are phylogenetically distinct species sharing a similar niche and swimming mode, suggesting that their analogous adipocyte/lipid distribution patterns are the consequence of convergent evolution. The potentially heterogeneous origin of adipose tissues has significant implications for the interpretation of their functional diversity.

Thermodynamic analysis of the squid mantle muscles and giant axon during slow swimming and jet escape propulsion

Squids have two substantially different types of muscle fibers: superficial mitochondria rich fibers, which perform aerobic respiration during slow swimming, and central mitochondria poor fibers, which perform anaerobic respiration during jet escape. A detailed thermodynamic analysis shows that during slow swimming, 3.82 J/(kg s) of chemical exergy is consumed, and a total muscle work of 0.28 J/(kg s) is produced. 0.27 J/(kg s) of this is produced by the fin to generate lift, and the rest by the mantle volume contraction. During the jet escape at a speed of 3 mantle length/s, squid consumes an exergy of 9.97 J/(kg s) and produces a muscle work of 0.16 J/(kg s). Exergy destruction rates during slow swimming and jet escape modes are 3.54 and 9.81 J/(kg s), respectively. Exergy destroyed because of the action potential propagation in the squid giant axon is calculated as 0.03 and 0.10 J/(kg s) for the slow and fast swimming modes, respectively.

Swimming patterns of the quadriflagellate Tetraflagellochloris mauritanica (Chlamydomonadales, Chlorophyceae).

Chlamydomonadales are elective subjects for the investigation of the problems related to locomotion and transport in biological fluid dynamics, whose resolution could enhance searching efficiency and assist in the avoidance of dangerous environments. In this paper, we elucidate the swimming behavior of Tetraflagellochloris mauritanica, a unicellular-multicellular alga belonging to the order Chlamydomonadales. This quadriflagellate alga has a complex swimming motion consisting of alternating swimming phases connected by in-place random reorientations and resting phases. It is capable of both forward and backward swimming, both being normal modes of swimming. The complex swimming behavior resembles the run-and-tumble motion of peritrichous bacteria, with in-place reorientation taking the place of tumbles. In the forward swimming, T.mauritanica shows a very efficient flagellar beat, with undulatory retrograde waves that run along the flagella to their tip. In the backward swimming, the flagella show a nonstereotypical synchronization mode, with a pattern that does not fit any of the modes present in the other Chlamydomonadales so far investigated.

Re-evaluation of batoid pectoral morphology reveals novel patterns of diversity among major lineages.

Batoids (Chondrichthyes: Batoidea) are a diverse group of cartilaginous fishes which comprise a monophyletic sister lineage to all neoselachians or modern sharks. All species in this group possess anteroposteriorly expanded-pectoral fins, giving them a unique disc-like body form. Reliance on pectoral fins for propulsion ranges from minimal (sawfish) to almost complete dependence (skates and rays). A recent study on the diversity of planform pectoral fin shape in batoids compared overall patterns of morphological variation within the group. However, inconsistent pectoral homology prevented the study from accurately representing relationships within and among major batoid taxa. With previous work in mind, we undertook an independent investigation of pectoral form in batoids and evaluated the implications of shape diversity on locomotion and lifestyle, particularly in the skates (Rajoidei) and rays (Myliobatoidei). We used geometric morphometrics with sliding semilandmarks to analyze pectoral fin outlines and also calculate fin aspect ratios (AR), a functional trait linked to locomotion. In agreement with previous work, our results indicated that much of the evolution of batoid pectoral shape has occurred along a morphological axis that is closely related to AR. For species where kinematic data were available, both shape and AR were associated with swimming mode. This work further revealed novel patterns of shape variation among batoids, including strong bimodality of shape in rays, an intermediate location of skate species in the morphospace between benthic/demersal and pelagic rays, and approximately parallel shape trajectories in the benthic/demersal rays and skates. Finally, manipulation of landmarks verified the need for a consistent and accurate definition of homology for the outcome and efficacy of analyses of pectoral form and function in batoids.

Methods matter: considering locomotory mode and respirometry technique when estimating metabolic rates of fishes.

Respirometry is frequently used to estimate metabolic rates and examine organismal responses to environmental change. Although a range of methodologies exists, it remains unclear whether differences in chamber design and exercise (type and duration) produce comparable results within individuals and whether the most appropriate method differs across taxa. We used a repeated-measures design to compare estimates of maximal and standard metabolic rates (MMR and SMR) in four coral reef fish species using the following three methods: (i) prolonged swimming in a traditional swimming respirometer; (ii) short-duration exhaustive chase with air exposure followed by resting respirometry; and (iii) short-duration exhaustive swimming in a circular chamber. We chose species that are steady/prolonged swimmers, using either a body-caudal fin or a median-paired fin swimming mode during routine swimming. Individual MMR estimates differed significantly depending on the method used. Swimming respirometry consistently provided the best (i.e. highest) estimate of MMR in all four species irrespective of swimming mode. Both short-duration protocols (exhaustive chase and swimming in a circular chamber) produced similar MMR estimates, which were up to 38% lower than those obtained during prolonged swimming. Furthermore, underestimates were not consistent across swimming modes or species, indicating that a general correction factor cannot be used. However, SMR estimates (upon recovery from both of the exhausting swimming methods) were consistent across both short-duration methods. Given the increasing use of metabolic data to assess organismal responses to environmental stressors, we recommend carefully considering respirometry protocols before experimentation. Specifically, results should not readily be compared across methods; discrepancies could result in misinterpretation of MMR and aerobic scope.

Aquatic locomotion of the terrestrial opossum Didelphis aurita (Didelphimorphia, Didelphidae) using undulatory swimming mode

AbstractThis note reports the first record of undulatory swimming mode by the black-eared opossum

Physostomous channel catfish, Ictalurus punctatus, modify swimming mode and buoyancy based on flow conditions.

The employment of gliding in aquatic animals as a means of conserving energy has been theoretically predicted and discussed for decades. Several studies have shown that some species glide, whereas others do not. Freshwater fish species that widely inhabit both lentic and lotic environments are thought to be able to adapt to fluctuating flow conditions in terms of locomotion. In adapting to the different functional demands of lentic and lotic environments on fish energetics, physostomous (open swim bladder) fish may optimise their locomotion and activity by controlling their net buoyancy; however, few buoyancy studies have been conducted on physostomous fish in the wild. We deployed accelerometers on free-ranging channel catfish, Ictalurus punctatus, in both lentic and lotic environments to quantify their swimming activity, and to determine their buoyancy condition preferences and whether gliding conserves energy. Individual comparisons of swimming efforts between ascent and descent phases revealed that all fish in the lentic environment had negative buoyancy. However, all individuals showed many descents without gliding phases, which was contrary to the behaviour predicted to minimise the cost of transport. The fact that significantly fewer gliding phases were observed in the lotic environment, together with the existence of neutrally buoyant fish, indicated that channel catfish seem to optimise their locomotion through buoyancy control based on flow conditions. The buoyancy optimisation of channel catfish relative to the flow conditions that they inhabit not only reflects differences in swimming behaviour but also provides new insights into the adaptation of physostome fish species to various freshwater environments.

Batoid locomotion: effects of speed on pectoral fin deformation in the little skate, Leucoraja erinacea.

Most batoids have a unique swimming mode in which thrust is generated by either oscillating or undulating expanded pectoral fins that form a disc. Only one previous study of the freshwater stingray has quantified three-dimensional motions of the wing, and no comparable data are available for marine batoid species that may differ considerably in their mode of locomotion. Here, we investigate three-dimensional kinematics of the pectoral wing of the little skate, Leucoraja erinacea, swimming steadily at two speeds [1 and 2bodylengths(BL)s-1]. We measured the motion of nine points in three dimensions during wing oscillation and determined that there are significant differences in movement amplitude among wing locations, as well as significant differences as speed increases in body angle, wing beat frequency and speed of the traveling wave on the wing. In addition, we analyzed differences in wing curvature with swimming speed. At 1 BL s-1, the pectoral wing is convex in shape during the downstroke along the medio-lateral fin midline, but at 2 BL s-1 the pectoral fin at this location cups into the flow, indicating active curvature control and fin stiffening. Wing kinematics of the little skate differed considerably from previous work on the freshwater stingray, which does not show active cupping of the whole fin on the downstroke.

Shape matters: Near-field fluid mechanics dominate the collective motions of ellipsoidal squirmers.

Microswimmers show a variety of collective motions. Despite extensive study, questions remain regarding the role of near-field fluid mechanics in collective motion. In this paper, we describe precisely the Stokes flow around hydrodynamically interacting ellipsoidal squirmers in a monolayer suspension. The results showed that various collective motions, such as ordering, aggregation, and whirls, are dominated by the swimming mode and the aspect ratio. The collective motions are mainly induced by near-field fluid mechanics, despite Stokes flow propagation over a long range. These results emphasize the importance of particle shape in collective motion.

Different ossification patterns of intermuscular bones in fish with different swimming modes.

ABSTRACTIntermuscular bones are found in the myosepta in teleosts. However, there is very little information on the development and ossification of these intermuscular bones. In this study, we performed an in-depth investigation of the ossification process during development in zebrafish (Danio rerio) and Japanese eel (Anguilla japonica). In Japanese eel, a typical anguilliform swimmer, the intermuscular bones ossified predominantly from the anterior to the posterior. By contrast, in the zebrafish, a sub-carangiform or carangiform swimmer, the intermuscular bones ossified predominantly from the posterior to the anterior regions of the fish. Furthermore, tail amputation affected the ossification of the intermuscular bones. The length of the intermuscular bones in the posterior area became significantly shorter in tail-amputated zebrafish and Japanese eels, and both had less active and lower swimming speeds; this indicates that swimming might induce the ossification of the intermuscular bones. Moreover, when a greater length of tail was amputated in the zebrafish, the intermuscular bones became even shorter. Tail amputation affected the length and ossification of intermuscular bones in the anterior part of the fish, close to the head, differently between the two fish: they became significantly shorter in the zebrafish, but did not in the Japanese eel. This might be because tail amputation did not significantly affect the undulations in the anterior of the Japanese eel, especially near the head. This study shows that the ossification of intermuscular bones might be induced through mechanical force loadings that are produced by swimming.

Two-dimensional slither swimming of sperm within a micrometre of a surface.

Sperm motion near surfaces plays a crucial role in fertilization, but the nature of this motion has not been resolved. Using total internal reflection fluorescence microscopy, we selectively imaged motile human and bull sperm located within one micron of a surface, revealing a distinct two-dimensional (2D) ‘slither' swimming mode whereby the full cell length (50–80 μm) is confined within 1 μm of a surface. This behaviour is distinct from bulk and near-wall swimming modes where the flagellar wave is helical and the head continuously rotates. The slither mode is intermittent (∼1 s, ∼70 μm), and in human sperm, is observed only for viscosities over 20 mPa·s. Bull sperm are slower in this surface-confined swimming mode, owing to a decrease in their flagellar wave amplitude. In contrast, human sperm are ∼50% faster—suggesting a strategy that is well suited to the highly viscous and confined lumen within the human fallopian tube.

Upstream range expansion by invasive round gobies: Is functional morphology important?

Range expansion by invasive taxa can be facilitated by a wide variety of life history traits, by behavioral repertoires, or via human-aided translocation. The round goby (Neogobius melanostomus) continues to expand its range in Europe and the Great Lakes watershed by moving upstream in tributary rivers and streams. It is not clear whether in-stream barriers (e.g., waterfalls, culverts, dams) will prevent further upstream expansion. Since other species within the Gobiidae are adept waterfall climbers, we investigated whether body scaling morphometrics, fused pelvic fin adhesion, or swimming patterns in round gobies were suggestive of climbing capability. Behaviorally, round gobies swam less frequently, limited their swimming mode to caudal thrusts, and sought shelter more as velocity increased. Attachment to the substrate via the fused pelvic fins was documented in a laboratory flume. Fish smaller than 5 cm or having a body mass less than 1.5 g had passive pelvic fin adhesion power sufficient to remain in place on a wetted, inclined plane, whereas larger fish did not. Small fish had significantly smaller pelvic fin area/length ratios relative to larger fish (P < 0.001), but significantly greater pelvic fin area/mass ratios (P < 0.001). Caudal fin area, but not fused pelvic fin area, exhibited an allometric growth pattern versus weight in large gobies, whereas neither caudal nor pelvic fin area showed allometric increases versus weight in small round gobies. Pelvic fin area in both large and small fish grew allometrically compared to length, whereas caudal fin area increased allometrically relative to length only in small fish. Fin morphometric results suggest, theoretically, that neither adults nor juveniles have pelvic fin areas large enough to support their weight against gravity forces, but the areas are sufficient to withstand hydraulic drag forces. Collectively, our results suggest that in-stream barriers which force fish to overcome gravity should be impassable, but that barriers allowing fish to remain submerged could be navigable.

Somersault of Paramecium in extremely confined environments.

We investigate various swimming modes of Paramecium in geometric confinements and a non-swimming self-bending behavior like a somersault, which is quite different from the previously reported behaviors. We observe that Paramecia execute directional sinusoidal trajectories in thick fluid films, whereas Paramecia meander around a localized region and execute frequent turns due to collisions with adjacent walls in thin fluid films. When Paramecia are further constrained in rectangular channels narrower than the length of the cell body, a fraction of meandering Paramecia buckle their body by pushing on the channel walls. The bucking (self-bending) of the cell body allows the Paramecium to reorient its anterior end and explore a completely new direction in extremely confined spaces. Using force deflection method, we quantify the Young’s modulus of the cell and estimate the swimming and bending powers exerted by Paramecium. The analysis shows that Paramecia can utilize a fraction of its swimming power to execute the self-bending maneuver within the confined channel and no extra power may be required for this new kind of self-bending behavior. This investigation sheds light on how micro-organisms can use the flexibility of the body to actively navigate within confined spaces.

Applying central pattern generators to control the robofish with oscillating pectoral fins

Purpose– This paper aims to develop a robofish with oscillating pectoral fins, and control it to mimic the bionic prototype by central pattern generators (CPGs).Design/methodology/approach– First, the oscillation characteristics of the cownose ray were analyzed quantitatively. Second, a robofish with multi-joint pectoral fins was developed according to the bionic morphology and kinematics. Third, the improved phase oscillator was established, which contains a spatial asymmetric coefficient and a temporal asymmetric coefficient. Moreover, the CPG network is created to mimic the cownose ray and accomplish three-dimensional (3D) motions. Finally, the experiments were done to test the authors ' works.Findings– The results demonstrate that the CPGs is effective to control the robofish to imitate the cownose ray realistically. In addition, the robofish is able to accomplish 3D motions of high maneuverability, and change among different swimming modes quickly and smoothly.Originality/value– The research provides the method to develop a robofish from both 3D morphology and kinematics. The motion analysis and CPG control make sure that the robofish has the features of high maneuverability and camouflage. It is useful for military underwater applications and underwater detections in narrow environments. Second, this work lays the foundation for the autonomous 3D control. Moreover, the robotic fish can be taken as a scientific tool for the fluid bionics research.

Quantitative flow analysis of swimming dynamics with coherent Lagrangian vortices.

Undulatory swimmers flex their bodies to displace water, and in turn, the flow feeds back into the dynamics of the swimmer. At moderate Reynolds number, the resulting flow structures are characterized by unsteady separation and alternating vortices in the wake. We use the flow field from simulations of a two-dimensional, incompressible viscous flow of an undulatory, self-propelled swimmer and detect the coherent Lagrangian vortices in the wake to dissect the driving momentum transfer mechanisms. The detected material vortex boundary encloses a Lagrangian control volume that serves to track back the vortex fluid and record its circulation and momentum history. We consider two swimming modes: the C-start escape and steady anguilliform swimming. The backward advection of the coherent Lagrangian vortices elucidates the geometry of the vorticity field and allows for monitoring the gain and decay of circulation and momentum transfer in the flow field. For steady swimming, momentum oscillations of the fish can largely be attributed to the momentum exchange with the vortex fluid. For the C-start, an additionally defined jet fluid region turns out to balance the high momentum change of the fish during the rapid start.

Influence of environmental conditions on foraging behaviour and its consequences on reproductive performance in little penguins

During the breeding season, seabirds are central place foragers and have to adapt their foraging behaviour in response to environmental variation to maximize efficiency and reproductive output. Due to its small size and swimming mode of transport, the little penguin (Eudyptula minor) is expected to be greatly susceptible to such fluctuations. The links between local-, meso- and macro-scale environmental conditions and inter-annual variation in foraging behaviour and reproductive performance of little penguins were investigated during three consecutive breeding seasons at two colonies in south-eastern Australia marked by contrasting oceanographic conditions. At a local scale, foraging effort was correlated positively with wind direction and negatively with wave height. At a regional scale, foraging effort of individuals from both colonies was negatively correlated with higher sea surface temperature (SST) off the Bonney Coast in the previous Austral summer, suggesting a weaker Bonney Upwelling event and a cascade of effects throughout the Bass Strait region. At a larger scale, the El Nino Southern Oscillation was also found to correlate with foraging behaviour, with lower foraging effort being observed during La Nina event. Although individuals increased their foraging effort during years with poorer conditions, they were not able to maintain high breeding success. In addition, peak egg-laying was found to coincide with a decrease in local SST and a peak of sea surface chlorophyll-a concentration. In conclusion, these results highlight how different environmental conditions could influence foraging behaviour and ultimately reproductive success of little penguins. It also showed that under certain circumstances, these individual strategies were not sufficient to cope with environmental variability.