Respirometry and swimming dynamics of the giant australian cuttlefish,sepia apama (mollusca,cephalopoda)

Swimming dynamics of the giant Australian cuttlefish, Sepia apama, were investigated using swimtunnel respirometry. Relationships between jet pressure, fin frequency, swimming speed and oxygen consumption were defined. Laboratory calibration of swimming parameters is necessary to allow estimates of swimming costs in the field.

Jet pressure was the best predictor of oxygen consumption with an averaged equation of MO₂?=?722 (jet pressure)?+?107?r ²?=?0.51. Individually, fin frequency and jet pressure correlated highly to swimming speed, but due to the complicated usage of finning and jetting, the correlation between swimming speed and oxygen consumption was weaker. Cuttlefish were not optimal swimtunnel subjects and could not swim at high speeds for extended periods. At 15°C and a swimming speed of 0.06?m?s^?1, the gross cost of transport was calculated to be 10.1?kg^?1?m?^?1, with a net cost of 4.1?kg^?1?m^?1.     

The locomotory function of the fins in the squid Loligo pealei

Kinematic data of high spatial and temporal resolution, acquired from image sequences of adult long-finned squid, Loligo pealei, during steady swimming in a flume, were used to examine the role of fins and the coordination between fin and jet propulsion in squid locomotion. Fin shape and body outlines were digitized and used to calculate fin wave speed, amplitude, frequency, angle of attack, body deformation, speed, and acceleration. L. pealei were observed to have two fin gait patterns with a transition at 1.4-1.8 mantle lengths per second (Lm s-1) marked by alternation between the two patterns. Fin motion in L. pealei exhibited characteristics of both traveling waves and flapping wings. At low speeds, fin motion was more wave-like; at high speeds, fin motion was more flap-like and was marked by regular periods during which the fins were wrapped tightly against the mantle. Fin cycle frequencies were dependent on swimming speed and gait, and obvious coordination between the fins and jet were observed. Fin wave speed, angle of attack, and body acceleration confirmed the role of fins in thrust production and revealed a role of fins at all swimming speeds by a transition from drag-based to lift-based thrust when fin wave speed dropped below swimming speed. Estimates of peak fin thrust were as high as 0.44-0.96 times peak jet thrust in steady swimming over the range of swimming speeds observed. Fin downstrokes generally contributed more to thrust than did upstrokes, especially at high speeds.     

The locomotory function of the fins in the squid Loligo pealei

Kinematic data of high spatial and temporal resolution, acquired from image sequences of adult long-finned squid, Loligo pealei, during steady swimming in a flume, were used to examine the role of fins and the coordination between fin and jet propulsion in squid locomotion. Fin shape and body outlines were digitized and used to calculate fin wave speed, amplitude, frequency, angle of attack, body deformation, speed, and acceleration. L. pealei were observed to have two fin gait patterns with a transition at 1.4-1.8 mantle lengths per second (L_m s^-1) marked by alternation between the two patterns. Fin motion in L. pealei exhibited characteristics of both traveling waves and flapping wings. At low speeds, fin motion was more wave-like; at high speeds, fin motion was more flap-like and was marked by regular periods during which the fins were wrapped tightly against the mantle. Fin cycle frequencies were dependent on swimming speed and gait, and obvious coordination between the fins and jet were observed. Fin wave speed, angle of attack, and body acceleration confirmed the role of fins in thrust production and revealed a role of fins at all swimming speeds by a transition from drag-based to lift-based thrust when fin wave speed dropped below swimming speed. Estimates of peak fin thrust were as high as 0.44-0.96 times peak jet thrust in steady swimming over the range of swimming speeds observed. Fin downstrokes generally contributed more to thrust than did upstrokes, especially at high speeds.     

Respirometry and swimming dynamics of the giant australian cuttlefish, sepia apama (mollusca, cephalopoda)

Swimming dynamics of the giant Australian cuttlefish, Sepia apama, were investigated using swimtunnel respirometry. Relationships between jet pressure, fin frequency, swimming speed and oxygen consumption were defined. Laboratory calibration of swimming parameters is necessary to allow estimates of swimming costs in the field.

Jet pressure was the best predictor of oxygen consumption with an averaged equation of MO₂ = 722 (jet pressure) + 107 r² = 0.51. Individually, fin frequency and jet pressure correlated highly to swimming speed, but due to the complicated usage of finning and jetting, the correlation between swimming speed and oxygen consumption was weaker. Cuttlefish were not optimal swimtunnel subjects and could not swim at high speeds for extended periods. At 15°C and a swimming speed of 0.06 m s^-1, the gross cost of transport was calculated to be 10.1 kg^-1 m ^-1, with a net cost of 4.1 kg^-1 m^-1.     

Propulsion Modeling and Analysis of a Biomimetic Swimmer

<正> We have studied a biomimetic swimmer based on the motion of bacteria such as Escherichia coli (E. coli) theoretically andexperimentally. The swimmer has an ellipsoidal cell body propelled by a helical filament. The performance of this swimmer wasestimated by modeling the dynamics of a swimmer in viscous fluid. We applied the Resistive Force Theory (RFT) on this modelto calculate the linear swimming speed and the efficiency of the model. A parametric study on linear velocity and efficiency tooptimize the design of this swimmer was demonstrated. In order to validate the theoretical results, a biomimetic swimmer wasfabricated and an experiment setup was prepared to measure the swimming speed and thrust force in silicone oil. The experimentalresults agree well with the theoretical values predicted by RFT. In addition, we studied the flow patterns surrounding thefilament with a finite element simulation with different Reynolds number (Re) to understand the mechanism of propulsion. Thesimulation results provide information on the nature of flow patterns generated by swimming filament. Furthermore, the thrustforces from the simulation were compared with the thrust forces from theory. The simulation results are in good agreement withthe theoretical results.     

Jet propulsion in salps (Tunicata: Thaliacea)

This paper describes the locomotion of salps by jet propulsion, from a combination of measurements of chamber pressures, static thrust, and electromyographic activity, with kinematic records of free-swimming and tethered salps. From such measurements, estimates are made of the thrust exerted, the drag incurred, and the work performed by single salps, and by chains of linked individuals. It is concluded that salp jet propulsion is a more economical process than is jet propulsion in other animals.     

Parametric Study of an Underwater Finned Propulsor Inspired by Bluespotted Ray

The performance of bluespotted rays was emulated in the design of a bioinspired underwater propulsor in the present work.First,the movement of a live bluespotted ray was captured for the swimming mode and useful information to the biomimetic mechanism design.By virtue of the modular and reeonfigurable design concept,an undulatory fin propulsion prototype was developed.With a proper experimental set-up,orthogonal experiments were conducted to investigate the effect of various fin design parameters on the propulsion speed,thrust,and power of the fish robot.The controllable fin parameters include frequency,amplitude,wavelength,fm shape,and undulatory mode.The significance of these parameters was also determined by using the variance analysis.The results demonstrate that the designed propulsor,imitating bluespotted rays with large expanded undulatory fins,is able to propel itself by changing various kinematic parameters.     

Strouhal number for flying and swimming in rhinoceros auklets Cerorhinca monocerata

Alcids propel themselves by flapping wings in air and water that have vastly different densities. We hypothesized that alcids change wing kinematics and maintain Strouhal numbers (St = fA/U, where f is wingbeat frequency, A is the wingbeat amplitude, and U is forward speed) within a certain range, to achieve efficient locomotion during both flying and swimming. We used acceleration and GPS loggers to measure the wingbeat frequency and forward speed of free‐ranging rhinoceros auklets Cerorhinca monocerata during both flying and swimming. We also measured wingbeat amplitude from video footage taken in the wild. On average, wingbeat frequency, forward speed, and wingbeat amplitude were 8.9 Hz, 15.3 m s^?1, and 0.39 m, respectively, during flying, and 2.6 Hz, 1.3 m s^?1, and 0.18 m, respectively, during swimming. The smaller wingbeat amplitude during swimming was achieved by partially folding the wings, while maintaining the dorso‐ventral wingbeat angle. Mean St was 0.23 during flying and 0.36 during swimming. The higher St value for swimming might be related to the higher thrust force required for propulsion in water. Our results suggest that rhinoceros auklets maintain St for both flying and swimming within the range (0.2–0.4) that propulsive efficiency is known to be high and St in both flying specialists and swimming specialists are known to converge.     

Ontogeny of squid mantle function: changes in the mechanics of escape-jet locomotion in the oval squid,Sepioteuthis lessoniana lesson, 1830

In Sepioteuthis lessoniana, the oval squid, ontogenetic changes in the kinematics of the mantle during escape-jet locomotion imply a decline in the relative mass flux of the escape jet and may affect the peak weight-specific thrust of the escape jet. To examine the relationship between ontogenetic changes in the kinematics of the mantle and the thrust generated during the escape jet, we simultaneously measured the peak thrust and the kinematics of the mantle of squid tethered to a force transducer. We tested an ontogenetic series of S. lessoniana that ranged in size from 5 to 40 mm dorsal mantle length (DML). In newly hatched squids, thrust peaked 40 ms after the start of the escape jet and reached a maximum of between 0.10 mN and 0.80 mN. In the largest animals, thrust peaked 70 ms after the start of the escape jet and reached a maximum of between 18 mN and 110 mN. Peak thrust was normalized by the wet weight of the squid and also by the cross-sectional area of the circumferential muscle that provides power for the escape jet. The weight-specific peak thrust of the escape jet averaged 0.36 in newly hatched squid and increased significantly to an average of 1.5 in the largest squids measured (P < 0.01). The thrust per unit area of circumferential muscle averaged 0.25 mN/mm(2) in hatchlings and increased significantly to an average of 1.4 mN/mm(2) in the largest animals tested (P < 0.01). The impulse of the escape jet was also lowest in newly hatched individuals (1.3 mN. s) and increased significantly to 1000 mN. s in the largest squids measured (P < 0.01). These ontogenetic changes in the mechanics of the escape jet suggest (1) that propulsion efficiency of the exhalant phase of the jet is highest in hatchlings, and (2) that the mechanics of the circumferential muscles of the mantle change during growth.     

Role of aerobic and anaerobic circular mantle muscle fibers in swimming squid: electromyography

Circular mantle muscle of squids and cuttlefishes consists of distinct zones of aerobic and anaerobic muscle fibers that are thought to have functional roles analogous to red and white muscle in fishes. To test predictions of the functional role of the circular muscle zones during swimming, electromyograms (EMGs) in conjunction with video footage were recorded from brief squid Lolliguncula brevis (5.0-6.8 cm dorsal mantle length, 10.9-18.3 g) swimming in a flume at speeds of 3-27 cm s(-1). In one set of experiments, in which EMGs were recorded from electrodes intersecting both the central anaerobic and peripheral aerobic circular mantle muscles, electrical activity was detected during each mantle contraction at all swimming speeds, and the amplitude and frequency of responses increased with speed. In another set of experiments, in which EMGs were recorded from electrodes placed in the central anaerobic circular muscle fibers alone, electrical activity was not detected during mantle contraction until speeds of about 15 cm s(-1), when EMG activity was sporadic. At speeds greater than 15 cm s(-1), the frequency of central circular muscle activity subsequently increased with swimming speed until maximum speeds of 21-27 cm s(-1), when muscular activity coincided with the majority of mantle contractions. These results indicate that peripheral aerobic circular muscle is used for low, intermediate, and probably high speeds, whereas central anaerobic circular muscle is recruited at intermediate speeds and used progressively more with speed for powerful, unsteady jetting. This is significant because it suggests that there is specialization and efficient use of locomotive muscle in squids.     

Upstroke Thrust, Drag Effects, and Stroke-glide Cycles in Wing-propelled Swimming by Birds

A number of bird species swim underwater by wing propulsion.Both among and within species, thrust generated during the recoveryphase (upstroke) varies from almost none to more than duringthe power phase (downstroke). More uneven thrust and unsteadyspeed may increase swimming costs because of greater inertialwork to accelerate the body fuselage (head and trunk), especiallywhen buoyant resistance is high during descent. I investigatedthese effects by varying relative fuselage speed during upstrokevs. downstroke in a model for wing-propelled murres which descendat relatively constant mean speed. As buoyant resistance declinedwith depth, the model varied stroke frequency and glide durationto maintain constant mean descent speed, stroke duration, andwork per stroke. When mean fuselage speed during the upstrokewas only 18% of that during the downstroke, stroke frequencywas constant with no gliding, so that power output was unchangedthroughout descent. When mean upstroke speed of the fuselagewas raised to 40% and 73% of mean downstroke speed, stroke frequencydeclined and gliding increased, so that power output decreasedrapidly with increasing depth. Greater inertial work with moreunequal fuselage speeds was a minor contributor to differencesin swimming costs. Instead, lower speeds during upstrokes requiredhigher speeds during downstrokes to maintain the same mean speed,resulting in nonlinear increases in drag at greater fuselagespeeds during the power phase. When fuselage speed was relativelyhigher during upstrokes, lower net drag at the same mean speedincreased the ability to glide between strokes, thereby decreasingthe cost of swimming.     

Effect of an Artificial Caudal Fin on the Performance of a Biomimetic Fish Robot Propelled by Piezoelectric Actuators

This paper addresses the design of a biomimetic fish robot actuated by piezoeeramic actuators and the effect of artificial caudal fins on the fish robot＇s performance. The limited bending displacement produced by a lightweight piezocomposite actuator was amplified and transformed into a large tail beat motion by means of a linkage system. Caudal fins that mimic the shape of a mackerel fin were fabricated for the purpose of examining the effect of caudal fm characteristics on thrust production at an operating frequency range. The thickness distribution of a real mackerel＇s fin was measured and used to design artificial caudal fins. The thrust performance of the biomimetic fish robot propelled by fins of various thicknesses was examined in terms of the Strouhal number, the Froude number, the Reynolds number, and the power consumption. For the same fm area and aspect ratio, an artificial caudal fin with a distributed thickness shows the best forward speed and the least power consumption.     

Design and Experiments of a Robotic Fish Imitating Cow-Nosed Ray

<正> The cow-nosed ray is studied as natural sample of a flapping-foil robotic fish.Body structure, motion discipline, and dynamicfoil deformation of cow-nosed ray are analyzed.Based on the analysis results, a robotic fish imitating cow-nosed ray,named Robo-ray Ⅱ, mainly composed of soft body, flexible ribs and pneumatic artificial muscles, is developed.Structure andswimming morphology of the robotic prototype are as that of a normal cow-nosed ray in nature.Key propulsion parameters ofRobo-ray Ⅱ at normal conditions, including the St Number at linear swimming, thrust coefficient at towing are studied throughexperiments.The suitable driving parameters are confirmed considering the efficiency and swimming velocity.Swimmingvelocity of 0.16 m·s~(-1)'and thrust coefficient of 0.56 in maximum are achieved in experiments.     

Foreflipper propulsion in the California sea lion, Zalophus californianus

The family Otariidae comprises the only group of marine mammals that habitually use their pectoral appendages to generate propulsive forces during swimming. This method of propulsion was examined in the California sea lion ( Zalophus californianus ), a representative member of the family. High-speed films were taken as a sea lion swam against a water current generated inside a large flow channel. Thrust production was determined by examining the body's movement at various stages of the propulsive cycle. Sea lions generate thrust continuously throughout the stroke. Over its initial three-quarters, foreflippers act as hydrofoils creating forward thrust and lift as they move vertically through the water. Thrust production is greatest, however, near the end of the stroke, when flippers are used as paddles and are oriented broad side to the oncoming flow. The force generated by this three-phased system of propulsion is likely to be greater than that attainable by either an exclusively lift-based hydrofoil or drag-based paddling style of swimming.
The kinematic changes that enable sea lions to change speed were also investigated. Film records revealed that stroke amplitude became greater with speed, although total stroke duration remained essentially constant. Sea lions increase stroke frequency with velocity but large variations in the measured values suggest that changes in amplitude and flipper angle of attack are also important parameters for modulating swimming speed.     

Transitions from Drag-based to Lift-based Propulsion in Mammalian Swimming

The evolution of fully aquatic mammals from quadrupedal, terrestrialmammals was associated with changes in morphology and swimmingmode. Drag is minimized by streamlining body shape and appendages.Improvement in speed, thrust production and efficiency is accomplishedby a change of swimming mode. Terrestrial and semiaquatic mammalsemploy drag-based propulsion with paddling appendages, whereasfully aquatic mammals use lift-based propulsion with oscillatinghydrofoils. Aerobic efficiencies are low for drag-based swimming,but reach a maximum of 30% for lift-based propulsion. Propulsiveefficiency is over 80% for lift-based swimming while only 33%for paddling. In addition to swimming mode, the transition tohigh performance propulsion was associated with a shift fromsurface to submerged swimming providing a reduction in transportcosts. The evolution of aquatic mammals from terrestrial ancestorsrequired increased swimming performance with minimal compromiseto terrestrial movement. Examination of modern analogs to transitionalswimming stages suggests that only slight modification to theneuromotor pattern used for terrestrial locomotion is requiredto allow for a change to lift-based propulsion.     

Fuzzy Vorticity Control of a Biomimetic Robotic Fish Using a Flapping Lunate Tail

Vorticity control mechanisms for flapping foils play a guiding role in both biomimetic thrust research and modeling the forward locomotion of animals with wings, fins, or tails. In this paper, a thrust-producing flapping lunate tail is studied through force and power measurements in a water channel. Proper vorticity control methods for flapping tails are discussed based on the vorticity control parameters: the dimensionless transverse amplitude, Strouhal number, angle of attack, and phase angle. Field tests are conducted on a free-swimming biomimetic robotic fish that uses a flapping tail. The results show that active control of Strouhal number using fuzzy logic control methods can efficiently reduce power consumption of the robotic fish and high swimming speeds can be obtained. A maximum speed of 1.17 length specific speed is obtained experimentally under conditions of optimal vorticity control. The St of the flapping tail is controlled within the range of 0.4～0.5.     

Swimming behavior and morphometry of the file shell Limaria fragilis

Swimming has evolved in only a few orders of Bivalves. In this study, the behavior, morphometry, and mechanics of swimming in the file shell Limaria fragilis were characterized and compared to the better understood scallops. Absolute swimming speed (cm?sec^?1) increased with increasing shell height, although relative swimming speed (body lengths?sec^?1) did not covary with shell height. The increase in absolute swimming speed was due to an increase in the distance covered during each valve clap as clap distance (cm?clap^?1) also increased with shell height while clapping frequency (claps?sec^?1) did not covary with animal size. Limaria fragilis displayed a variety of morphological changes related to size. Shell length was negatively allometric with shell height indicating the shell became proportionately slimmer in larger animals. Dry shell mass was negatively allometric with shell height, while both dry adductor muscle mass and dry mantle + tentacle mass were positively allometric. Autotomy of mantle tentacles significantly decreased clap distance by 13% without affecting clapping frequency or swimming speed.     

Effect of leg kick on active drag in front-crawl swimming: Comparison of whole stroke and arms-only stroke during front-crawl and the streamlined position

The purpose of this study was to examine the effect of leg kick on the resistance force in front-crawl swimming. The active drag in front-crawl swimming with and without leg motion was evaluated using measured values of residual thrust (MRT method) and compared with the passive drag of the streamlined position (SP) for the same swimmers. Seven male competitive swimmers participated in this study, and the testing was conducted in a swimming flume. Each swimmer performed front-crawl under two conditions: using arms and legs (whole stroke: WS) and using arms only (arms-only stroke: AS). Active drag and passive drag were measured at swimming velocities of 1.1 and 1.3 m s⁻¹ using load cells connected to the swimmer via wires. We calculated a drag coefficient to compare the resistances of the WS, AS and SP at each velocity. For both the WS and AS at both swimming velocities, active drag coefficient was found to be about 1.6–1.9 times larger than that in passive conditions. In contrast, although leg movement did not cause a difference in drag coefficient for front-crawl swimming, there was a large effect size (d = 1.43) at 1.3 m s⁻¹. Therefore, although upper and lower limb movements increase resistance compared to the passive condition, the effect of leg kick on drag may depend on swimming velocity.     

Swimming Speed of Larval Snail Does Not Correlate with Size and Ciliary Beat Frequency

Many marine invertebrates have planktonic larvae with cilia used for both propulsion and capturing of food particles. Hence, changes in ciliary activity have implications for larval nutrition and ability to navigate the water column, which in turn affect survival and dispersal. Using high-speed high-resolution microvideography, we examined the relationship between swimming speed, velar arrangements, and ciliary beat frequency of freely swimming veliger larvae of the gastropod Crepidula fornicata over the course of larval development. Average swimming speed was greatest 6 days post hatching, suggesting a reduction in swimming speed towards settlement. At a given age, veliger larvae have highly variable speeds (0.8–4 body lengths s⁻¹) that are independent of shell size. Contrary to the hypothesis that an increase in ciliary beat frequency increases work done, and therefore speed, there was no significant correlation between swimming speed and ciliary beat frequency. Instead, there are significant correlations between swimming speed and visible area of the velar lobe, and distance between centroids of velum and larval shell. These observations suggest an alternative hypothesis that, instead of modifying ciliary beat frequency, larval C. fornicata modify swimming through adjustment of velum extension or orientation. The ability to adjust velum position could influence particle capture efficiency and fluid disturbance and help promote survival in the plankton.     

Reynolds number limits for jet propulsion: a numerical study of simplified jellyfish

The Scallop theorem states that reciprocal methods of locomotion, such as jet propulsion or paddling, will not work in Stokes flow (Reynolds number=0). In nature the effective limit of jet propulsion is still in the range where inertial forces are significant. It appears that almost all animals that use jet propulsion swim at Reynolds numbers (Re) of about 5 or more. Juvenile squid and octopods hatch from the egg already swimming in this inertial regime. Juvenile jellyfish, or ephyrae, break off from polyps swimming at Re greater than 5. Many other organisms, such as scallops, rarely swim at Re less than 100. The limitations of jet propulsion at intermediate Re is explored here using the immersed boundary method to solve the 2D Navier-Stokes equations coupled to the motion of a simplified jellyfish. The contraction and expansion kinematics are prescribed, but the forward and backward swimming motions of the idealized jellyfish are emergent properties determined by the resulting fluid dynamics. Simulations are performed for both an oblate bell shape using a paddling mode of swimming and a prolate bell shape using jet propulsion. Average forward velocities and work put into the system are calculated for Re between 1 and 320. The results show that forward velocities rapidly decay with decreasing Re for all bell shapes when Re<10. Similarly, the work required to generate the pulsing motion increases significantly for Re<10. When compared to actual organisms, the swimming velocities and vortex separation patterns for the model prolate agree with those observed in Nemopsis bachei. The forward swimming velocities of the model oblate jellyfish after two pulse cycles are comparable to those reported for Aurelia aurita, but discrepancies are observed in the vortex dynamics between when the 2D model oblate jellyfish and the organism. This discrepancy is likely due to a combination of the differences between the 3D reality of the jellyfish and the 2D simplification, as well as the rigidity of the time varying geometry imposed by the idealized model.