Tail Beat Amplitude Research Articles

Effect of caudal fin height on swimming kinematics in the mackerel Scomber scombrus L.

Geometry of the caudal fin of the mackerel Scomber scombrus was modified to examine the effect of the height of the fin on swimming kinematics. Mackerel with reduced caudal fin height had significantly shorter stride length and higher tail beat amplitude compared with the control having an intact caudal fin (P < 0·005).

THE MYSTERIOUS LITTLE FATTY FIN

Sitting on the back of many fishes, in between the dorsal fin and the tail,is an enigmatic little fatty flap of skin called the adipose fin. It looks a bit like an extra dorsal fin, and though it's present in eight large groups of fishes, no one knows why it's there. It might help prevent flow from wrapping over the top of the fish; it might help counteract forces from the anal fin,which is in about the same place, but on the ventral side; it might be a flow sensor; or it might not do anything, persisting due to developmental constraints. Whatever the fin's function, most fisheries scientists think it's not terribly important, because they regularly snip it off to mark millions of hatchery fish released into the wild each year.Thomas Reimchen and Nicola Temple at the University of Victoria in Canada devised a simple test to find out how important the adipose fin really is. They swam steelhead trout at speeds between about one and three body lengths per second, measured the tail beat frequency and amplitude, then clipped the adipose fin off and made the same measurements again. They expected that the standard fisheries wisdom would be right, and they'd see no difference between the clipped and unclipped fish.But, in fact, the fish with clipped fins tended to use a higher tail beat amplitude at all swimming speeds. It wasn't a lot higher - only about 8% on average - but it was usually a significant difference, except in the smallest fish. Reimchen and Temple worried, though, that the effect might not represent any intrinsic function of the adipose fin, but just the trauma of having a fin snipped off. So they tested another batch of fish in which they made a scratch along the base of the adipose fin, without actually cutting the fin off. The scratched fish swam the same as the unscratched ones, eliminating the trauma as a possible cause of the increase in tail beat amplitude.Why would snipping off the adipose fin lead to a higher amplitude? Reimchen and Temple can only speculate, but they raise some important questions. Perhaps the fin generates some thrust on its own, or makes vortices that increase the thrust of the tail fin. Without the extra thrust, trout would have to compensate by swimming harder. Or the adipose fin might help the fish swim more efficiently by sensing vortices upstream of the tail fin. Trout might counteract the lower efficiency after their adipose fin is clipped by using a higher tail beat amplitude. The suggestion that the adipose fin functions as a flow sensor seems plausible, since Reimchen found some small nerves running to the base of the fin.Whatever the mechanism, it appears that trout with clipped adipose fins must swim harder. It would be useful to compare the oxygen consumption of clipped and unclipped fish, to verify that swimming without an adipose fin is truly more difficult. But Reimchen and Temple's results should give fisheries scientists pause for thought, because they could have serious consequences for the millions of fish with clipped adipose fins released from hatcheries each year.

The Kármán gait: novel body kinematics of rainbow trout swimming in a vortex street

Most fishes commonly experience unsteady flows and hydrodynamic perturbations during their lifetime. In this study, we provide evidence that rainbow trout Oncorhynchus mykiss voluntarily alter their body kinematics when interacting with vortices present in the environment that are not self-generated. To demonstrate this, we measured axial swimming kinematics in response to changes in known hydrodynamic wake characteristics. We compared trout swimming in the Kármán street behind different diameter cylinders (2.5 and 5 cm) at two flow speeds (2.5 and 4.5 L s(-1), where L is total body length) to trout swimming in the free stream and in the cylinder bow wake. Trout swimming behind cylinders adopt a distinctive, previously undescribed pattern of movement in order to hold station, which we term the Kármán gait. During this gait, body amplitudes and curvatures are much larger than those of trout swimming at an equivalent flow velocity in the absence of a cylinder. Tail-beat frequency is not only lower than might be expected for a trout swimming in the reduced flow behind a cylinder, but also matches the vortex shedding frequency of the cylinder. Therefore, in addition to choosing to be in the slower flow velocity offered behind a cylinder (drafting), trout are also altering their body kinematics to synchronize with the shed vortices (tuning), using a mechanism that may not involve propulsive locomotion. This behavior is most distinctive when cylinder diameter is large relative to fish length. While tuning, trout have a longer body wavelength than the prescribed wake wavelength, indicating that only certain regions of the body may need to be oriented in a consistent manner to the oncoming vortices. Our results suggest that fish can capture energy from vortices generated by the environment to maintain station in downstream flow. Interestingly, trout swimming in front of a cylinder display lower tail-beat amplitudes and body wave speeds than trout subjected to any of the other treatments, implying that the bow wake may be the most energetically favorable region for a fish to hold station near a cylinder.

Ontogenetic changes of swimming kinematics in a semi-aquatic reptile (Crocodylus porosus)

Semi-aquatic animals represent a transitional locomotor condition characterised by the possession of morphological features that allow locomotion both in water and on land. Most ecologically important behaviours of crocodilians occur in the water, raising the question of whether their 'terrestrial construction' constrains aquatic locomotion. Moreover, the demands for aquatic locomotion change with life-history stage. It was the aim of this research to determine the kinematic characteristics and efficiency of aquatic locomotion in different-sized crocodiles (Crocodylus porosus). Aquatic propulsion was achieved primarily by tail undulations, and the use of limbs during swimming was observed only in very small animals or at low swimming velocities in larger animals. Over the range of swimming speeds we examined, tail beat amplitude did not change with increasing velocity, but amplitude increased significantly with body length. However, amplitude expressed relative to body length decreased with increasing body length. Tail beat frequency increased with swimming velocity but there were no differences in frequency between different-sized animals. Mechanical power generated during swimming and thrust increased non-linearly with swimming velocity, but disproportionally so that kinematic efficiency decreased with increasing swimming velocity. The importance of unsteady forces, expressed as the reduced frequency, increased with increasing swimming velocity. Amplitude is the main determinant of body-size-related increases in swimming velocity but, compared with aquatic mammals and fish, crocodiles are slow swimmers probably because of constraints imposed by muscle performance and unsteady forces opposing forward movement. Nonetheless, the kinematic efficiency of aquatic locomotion in crocodiles is comparable to that of fully aquatic mammals, and it is considerably greater than that of semi-aquatic mammals.

Swimming in needlefish (Belonidae): anguilliform locomotion with fins.

The Atlantic needlefish (Strongylura marina) is a unique anguilliform swimmer in that it possesses prominent fins, lives in coastal surface-waters, and can propel itself across the surface of the water to escape predators. In a laboratory flow tank, steadily swimming needlefish perform a speed-dependent suite of behaviors while maintaining at least a half wavelength of undulation on the body at all times. To investigate the effects of discrete fins on anguilliform swimming, I used high-speed video to record body and fin kinematics at swimming speeds ranging from 0.25 to 2.0 L s(-1) (where L is the total body length). Analysis of axial kinematics indicates that needlefish are less efficient anguilliform swimmers than eels, indicated by their lower slip values. Body amplitudes increase with swimming speed, but unlike most fishes, tail-beat amplitude increases linearly and does not plateau at maximal swimming speeds. At 2.0 L s(-1), the propulsive wave shortens and decelerates as it travels posteriorly, owing to the prominence of the median fins in the caudal region of the body. Analyses of fin kinematics show that at 1.0 L s(-1) the dorsal and anal fins are slightly less than 180 degrees out of phase with the body and approximately 225 degrees out of phase with the caudal fin. Needlefish exhibit two gait transitions using their pectoral fins. At 0.25 L s(-1), the pectoral fins oscillate but do not produce thrust, at 1.0 L s(-1) they are held abducted from the body, forming a positive dihedral that may reduce rolling moments, and above 2.0 L s(-1) they remain completely adducted.

Kinematics of plaice, Pleuronectes platessa, and cod, Gadus morhua, swimming near the bottom.

The kinematics of plaice (Pleuronectes platessa, L=22.1 cm) and cod (Gadus morhua, L=25.0 cm, where L is total fish length) swimming at various speeds at the bottom and lifted to heights, h, of 10, 50 and 100 mm by a thin-wire grid were measured. For cod, tailbeat frequency, amplitude, body and fin span and propulsive wavelength were unaffected by h and varied with speed as described for fusiform pelagic species. In contrast, the kinematics of plaice was affected by h. Body and fin spans and propulsive wavelength were independent of swimming speed and h. Tailbeat amplitude was independent of swimming speed, but averaged 1.5 cm at h=0 and 2.5 cm at h> or = 10 mm. Plaice tailbeat frequency increased with swimming speed for fish at the bottom but was independent of swimming speed at h=10, 50 and 100 mm, averaging 4.6, 6.0 and 5.8 Hz respectively. Total mechanical power, P, produced by propulsive movements calculated from the bulk-momentum form of elongated slender-body theory was similar for cod and plaice swimming at the bottom but, at h> or = 10 mm, P for plaice was larger than that for cod. Plaice support their weight in water by swimming at a small tilt angle. The small changes in swimming kinematics with swimming speed are attributed to decreasing induced power costs to support the weight as speed increases. The contribution of the tail to power output increased monotonically with the tail gap/span ratio, z/B, for z/B=0.23 (h=0 mm) to z/B=1.1 (h=50 mm). The smaller tailbeat amplitude of the tail decreased both z/B and the power output for plaice swimming at the bottom. For the maximum body and fin span of plaice, the contribution to power output increased for local z/B values of 0.044 (h-0 mm) to 0.1 (h=10 mm) and declined somewhat at larger values of z/B. The smaller effect of the bottom on power output of the large-span anterior body sections may result from the resorption of much of the upstream wake at the re-entrant downstream tail.

Does pregnancy affect swimming performance of female Mosquitofish, Gambusia affinis?

Summary1. The cost of reproduction due to limiting of the reproductive female’s locomotion capability has been suggested many times, but has rarely been directly examined, especially in fishes. Here, the effect of pregnancy on swimming performance in the viviparous Mosquitofish, Gambusia affinis, was studied.2. Eight females of G. affinis were isolated, each in a separate aquarium, and critical swimming speed (Ucrit), body mass (BM) and cross‐section area were measured every 5 days from the beginning of the pregnancy until 2–4 days after parturition.3. Swimming kinematics (tail beat frequency and amplitude) was measured in non‐pregnant and pregnant females at different swimming speeds.4. BM increased during pregnancy from 0·47 ± 0·13 g to 0·72 ± 0·19 g, and the cross‐section area also increased during pregnancy from 0·21 ± 0·06 cm2 to 0·32 ± 0·07 cm2. Ucrit decreased from 25·0 ± 1·3 cm s−1 before pregnancy to 20·1 ± 1·5 cm s−1 just before parturition, and returned to 24·7 ± 1·4 cm s−1 2–4 days after parturition. Interindividual variation was repeatable and reflects real differences among individuals.5. Swimming kinematics was not affected by pregnancy.6. The results suggest that reductions in Ucrit are probably because of aerobic constraints and not necessarily due to hydrodynamic changes resulting from changing in body form or plasticity. Moreover, the reduction in Ucrit is, potentially, a ‘cost of reproduction’ owing to decrease in the ability to gain food during pregnancy in G. affinis females.

Effects of temperature on sustained swimming performance and swimming kinematics of the chub mackerel Scomber japonicus.

The effects of a 6 degrees C difference in water temperature on maximum sustained swimming speed, swimming energetics and swimming kinematics were measured in the chub mackerel Scomber japonicus (Teleostei: Scombridae), a primarily coastal, pelagic predator that inhabits subtropical and temperate transition waters of the Atlantic, Pacific and Indian Oceans. New data for chub mackerel acclimated to 18 degrees C are compared with published data from our laboratory at 24 degrees C. Twelve individuals acclimated to each of two temperatures (15.6-26.3 cm fork length, FL, and 34-179 g at 18 degrees C; 14.0-24.7 cm FL and 26-156 g at 24 degrees C) swam at a range of speeds in a temperature-controlled Brett-type respirometer, at the respective acclimation temperature. At a given fish size, the maximum speed that S. japonicus was able to maintain for a 30-min period, while swimming steadily using slow, oxidative locomotor muscle (U(max,c)), was significantly greater at 24 than at 18 degrees C (52.5-97.5 cm s(-1) at 18 degrees C and 70-120 cm s(-1) at 24 degrees C). At a given speed and fish size, the rate of oxygen consumption (VO(2)) was significantly higher at 24 than at 18 degrees C because of a higher net cost of transport (1073-4617 J km(-1) kg(-1) at 18 degrees C and 2708-14895 J km(-1) kg(-1) at 24 degrees C). Standard metabolic rate, calculated by extrapolating the logO(2) versus swimming speed relationship to zero speed, did not vary significantly with temperature or fish mass (126.4+/-67.2 mg O(2) h(-1) kg(-1) at 18 degrees C and 143.2+/-80.3 mg O(2) h(-1) kg(-1) at 24 degrees C; means +/- S.D., N=12). Swimming kinematics was quantified from high-speed (120 Hz) video recordings analyzed with a computerized, two-dimensional motion-analysis system. At a given speed and fish size, there were no significant effects of temperature on tail-beat frequency, tail-beat amplitude or stride length, but propulsive wavelength increased significantly with temperature as a result of an increase in propulsive wave velocity. Thus, the main effects of temperature on chub mackerel swimming were increases in both U(max,c) and the net cost of swimming at 24 degrees C. Like other fishes, S. japonicus apparently must recruit more slow, oxidative muscle fibers to swim at a given sustainable speed at the lower temperature because of the reduced power output. Thus, the 24 degrees C mackerel reach a higher speed before they must recruit the fast, glycolytic fibers, thereby increasing U(max,c) at 24 degrees C. By quantifying in vivo the effects of temperature on the swimming performance of an ectothermic species that is closely related to the endothermic tunas, this study also provides evidence that maintaining the temperature of the slow, oxidative locomotor muscle at 6 degrees C or more above ambient water temperature in tunas should significantly increase sustainable swimming speeds, but also increase the energetic cost of swimming, unless cardiac output limits muscle performance.

Recovery of locomotion correlated with axonal regeneration after a complete spinal transection in the eel

This research has examined the relationship between axonal regeneration and the return of normal movement following complete transection of the spinal cord. We made measurements of tail beat frequency and amplitude of the caudal body wave from video recordings of eels ( Anguilla anguilla) swimming in a water tunnel at several speeds. Each eel was then anaesthetised and the spinal cord cut caudal to the anus; in some animals the resulting gap was filled with a rubber block. All animals were kept at 25°C for recovery periods ranging from 7 to 128 days, during which their swimming performance was monitored regularly. Each fish was then re-anaesthetised and perfused with fixative and the regrowing descending axons labelled with 1,1′-diotadecyl-3,3,3′,3′-tetramethylindocarbocyanine perchlorate. For all animals and at all speeds after surgery, tail beat frequency increased, while amplitude decreased. In non-blocked animals, an improvement in performance was first seen from 8 days following transection and thereafter tail beat frequency decreased progressively until it had returned to normal after 35 to 45 days, while amplitude remained below baseline until at least 45 days. In these animals, few axonal growth cones had penetrated the caudal stump by 7 days, but some had extended as much as 3 mm by 15 days. Many had reached as far as 6 mm between 25 and 36 days, while by 128 days they had progressed up to 10.5 mm. Contralateral crossing was never observed. Functional recovery was never witnessed in animals in which the cord had been blocked and these eels swam at all times with elevated tail beat frequency and reduced caudal amplitude. No labelled axons could be traced into the caudal spinal cord at any recovery stage in such animals. We conclude that re-innervation of only 1–2 segments caudal to the injury is necessary for functional recovery, although continued axonal growth may be important for the refinement of some aspects of movement.

Caudal differential pressure as a predictor of swimming speed of cod (Gadus morhua).

We report the results of an experiment designed to investigate the feasibility of using differential pressure to estimate the swimming speed and metabolic rate of Atlantic cod (Gadus morhua). Seven cod were fitted with a miniature differential pressure sensor mounted on one side of the caudal peduncle immediately anterior to the base of the caudal fin rays. Relationships between differential pressure, tailbeat frequency, tailbeat amplitude, swimming speed and rate of oxygen consumption ((O(2))) were determined as a function of the swimming speed of cod swimming at 5 degrees C in a recirculating 'Brett-style' respirometer. Tailbeat differential pressure, tailbeat amplitude and tailbeat frequency were highly correlated with swimming speed. The average or integrated pressure ranged from 0 to 150 Pa for speeds up to 0.8 m s(-1) (1.1 L s(-1), where L is total body length), while the 'pressure difference' (maximum minus minimum pressure) ranged from 0 to 900 Pa. Small changes in swimming speed of less than 0.05 m s(-1) were readily detected as differences in tailbeat pressure. Burst swimming in the respirometer resulted in huge pressure 'bursts' of up to 5000 Pa 'pressure difference'. The rate of oxygen consumption increased exponentially and was highly correlated with swimming speed (r(2)=0.77). The rate of oxygen consumption was also correlated with tailbeat integrated pressure (r(2)=0.68) and with differential pressure (r(2)=0.43); regression correlations were always greater for individuals than for combined data from all cod. The results detailed in this study indicate that an ultrasonic differential pressure transmitter would enable accurate estimates of the swimming speed, rates of oxygen consumption and activity patterns of free-ranging fish in nature.

Rainbow trout display a developmental shift in red muscle kinetics, swimming kinematics and myosin heavy chain isoform

Both activation and relaxation times of rainbow trout Oncorhynchus mykiss red muscle were shorter in parr than in older juveniles. Furthermore, parr red muscle had a faster maximum shortening velocity than that of older fish, as estimated with the force‐clamp technique. Parr swam with higher tailbeat frequencies and lower tailbeat amplitude than did older fish across a range of length‐specific steady swimming speeds. The developmental shift in contraction kinetics of red muscle and steady swimming kinematics was associated with a reduction from two or three myosin heavy chain isoforms in parr to one in older juveniles. This transition provides a mechanism to explain the variations in muscle contraction kinetics and swimming performance.

Tail kinematics of the chub mackerel Scomber japonicus: testing the homocercal tail model of fish propulsion.

Scombrid fishes possess a homocercal caudal fin with reduced intrinsic musculature and dorso-ventrally symmetrical external and internal morphology. Because of this symmetrical morphology, it has often been assumed that scombrid caudal fins function as predicted by the homocercal tail model. According to that model, the caudal fin moves in a dorso-ventrally symmetrical manner and produces no vertical lift during steady swimming. To test this hypothesis, we examined the tail kinematics of chub mackerel, Scomber japonicus (24.8+/-1.3 cm total length, L). Markers were placed on the caudal fin to identify specific regions of the tail, and swimming chub mackerel were videotaped from lateral and posterior views, allowing a three-dimensional analysis of tail motion. Analysis of tail kinematics suggests that, at a range of swimming speeds (1.2-3.0 L s(-)(1)), the dorsal lobe of the tail undergoes a 15 % greater lateral excursion than does the ventral lobe. Lateral excursion of the dorsal tail-tip also increases significantly by 32 % over this range of speeds, indicating a substantial increase in tail-beat amplitude with speed. In addition, if the tail were functioning in a dorso-ventrally symmetrical manner, the tail should subtend an angle of 90 degrees relative to the frontal (or xz) plane throughout the tail beat. Three-dimensional kinematic analyses reveal that the caudal fin actually reaches a minimum xz angle of 79.8 degrees. In addition, there is no difference between the angle subtended by the caudal peduncle (which is anterior to the intrinsic tail musculature) and that subtended by the posterior lobes of the tail. Thus, asymmetrical movements of the tail are apparently generated by the axial musculature and transmitted posteriorly to the caudal fin. These results suggest that the caudal fin of the chub mackerel is not functioning symmetrically according to the homocercal model and could produce upward lift during steady swimming.

Going against the flow: an examination of the propulsive movements made by young brook trout in streams

We examined the propulsive movements and behaviour of young-of-the-year (YOY) brook trout (Salvelinus fontinalis) swimming in their natal streams. Our findings demonstrated that swimming performance was influenced by temporal and spatial heterogeneity in water flow. Pectoral fins were used commonly, even by individuals swimming in fast flowing water. There also was spatial variation in the speed attained for a given tail-beat frequency and amplitude. After controlling statistically for variation in spatial location, fork length, and tail-beat amplitude, the swimming speeds brook trout attained for a given tail-beat frequency were lower than values expected from laboratory studies of steady swimming but higher than values expected from laboratory studies of unsteady swimming in standing water. Trout holding station made short-term adjustments in tail-beat frequency also suggesting a degree of unsteady swimming. A field experiment demonstrated that introduction of a current-velocity refuge reduced swimming costs by 10%, on average, without affecting the frequency of foraging attempts made.

Fish foot prints: morphology and energetics of the wake behind a continuously swimming mullet (Chelon labrosus Risso).

The structure of the wake behind a continuously swimming mullet was analysed qualitatively and quantitatively by applying two-dimensional particle image velocimetry. A detailed analysis of the flow pattern and of the swimming movements of the fish allowed us to derive a kinematic explanation of the flow pattern as well as an estimate of the relative contributions of the body and the tail to thrust production. During active propulsion, the undulatory swimming fish shed a wake consisting in the medio-frontal plane of a rearward, zigzagging jet flow between alternating vortices. The fish shed one vortex per half tailbeat when the tail reached its most lateral position. Part of the circulation shed in the vortices had been generated previously on the body by the transverse body wave travelling down the body. This undulatory pump mechanism accounted for less than half of the energy shed in the wake. The remainder was generated by the tail. The vortex spacing matched the tailbeat amplitude and the stride length.

Anguilliform locomotion in an elongate salamander (Siren intermedia): effects of speed on axial undulatory movements

Many workers interested in the mechanics and kinematics of undulatory aquatic locomotion have examined swimming in fishes that use a carangiform or subcarangiform mode. Few empirical data exist describing and quantifying the movements of elongate animals using an anguilliform mode of swimming. Using high-speed video, I examine the axial undulatory kinematics of an elongate salamander, Siren intermedia, in order to provide data on how patterns of movement during swimming vary with body position and swimming speed. In addition, swimming kinematics are compared with those of other elongate vertebrates to assess the similarity of undulatory movements within the anguilliform locomotor mode. In Siren, most kinematic patterns vary with longitudinal position. Tailbeat period and frequency, stride length, Froude efficiency and the lateral velocity and angle of attack of tail segments all vary significantly with swimming speed. Although swimming speed does not show a statistically significant effect on kinematic variables such as maximum undulatory amplitude (which increases non-linearly along the body), intervertebral flexion and path angle, examination of the data suggests that speed probably has subtle and site-specific effects on these variables which are not detected here owing to the small sample size. Maximum lateral displacement and flexion do not coincide in time within a given tailbeat cycle. Furthermore, the maximum orientation (angle with respect to the animal's direction of forward movement) and lateral velocity of tail segments also do not coincide in time. Comparison of undulatory movements among diverse anguilliform swimmers suggests substantial variation across taxa in parameters such as tailbeat amplitude and in the relationship between tailbeat frequency and swimming speed. This variation is probably due, in part, to external morphological differences in the shape of the trunk and tail among these taxa.

Kinematics and critical swimming speed of juvenile scalloped hammerhead sharks

Kinematics and critical swimming speed (Ucrit) of juvenile scalloped hammerhead sharks Sphyrna lewini were measured in a Brett-type flume (635 l). Kinematic parameters were also measured in sharks swimming in a large pond for comparison with those of sharks swimming in the flume. Sharks in the flume exhibited a mean Ucrit of 65±11 cm s-1 (± s.d.) or 1.17±0.21 body lengths per second (L s-1), which are similar to values for other species of sharks. In both the flume and pond, tailbeat frequency (TBF) and stride length (LS) increased linearly with increases in relative swimming speed (Urel=body lengths traveled per second). In the flume, tailbeat amplitude (TBA) decreased with increasing speed whereas TBA did not change with speed in the pond. Differences in TBF and LS between sharks swimming in the flume and the pond decreased with increases in Urel. Sharks swimming at slow speeds (e.g. 0.55 L s-1) in the pond had LS 19 % longer and TBF 21 % lower than sharks in the flume at the same Urel. This implies that sharks in the flume expended more energy while swimming at comparable velocities. Comparative measurements of swimming kinematics from sharks in the pond can be used to correct for effects of the flume on shark swimming kinematics and energetics.

Functions of fish skin: flexural stiffness and steady swimming of longnose gar, Lepisosteus osseus

The functions of fish skin during swimming remain enigmatic. Does skin stiffen the body and alter the propagation of the axial undulatory wave? To address this question, we measured the skin's in situ flexural stiffness and in vivo mechanical role in the longnose gar Lepisosteus osseus. To measure flexural stiffness, dead gar were gripped and bent in a device that measured applied bending moment (N m) and the resulting midline curvature (m-1). From these values, the flexural stiffness of the body (EI in N m2) was calculated before and after sequential alterations of skin structure. Cutting of the dermis between two caudal scale rows significantly reduced the flexural stiffness of the body and increased the neutral zone of curvature, a region of bending without detectable stiffness. Neither bending property was significantly altered by the removal of a caudal scale row. These alterations in skin structure were also made in live gar and the kinematics of steady swimming was measured before and after each treatment. Cutting of the dermis between two caudal scale rows, performed under anesthesia, changed the swimming kinematics of the fish: tailbeat frequency (Hz) and propulsive wave speed (body lengths per second, L s-1) decreased, while the depth (in L) of the trailing edge of the tail increased. The decreases in tailbeat frequency and wave speed are consistent with predictions of the theory of forced, harmonic vibrations; wave speed, if equated with resonance frequency, is proportional to the square root of a structure's stiffness. While it did not significantly reduce the body's flexural stiffness, surgical removal of a caudal scale row resulted in increased tailbeat amplitude and the relative total hydrodynamic power. In an attempt to understand the specific function of the scale row, we propose a model in which a scale row resists medio-lateral force applied by a single myomere, thus functioning to enhance mechanical advantage for bending. Finally, surgical removal of a precaudal scale row did not significantly alter any of the kinematic variables. This lack of effect is associated with a lower midline curvature of the precaudal region during swimming compared with that of the caudal region. Overall, these results demonstrate a causal relationship between skin, the passive flexural stiffness it imparts to the body and the influence of body stiffness on the undulatory wave speed and cycle frequency at which gar choose to swim.

Mechanical control of swimming speed: stiffness and axial wave form in undulating fish models

The purpose of this study was to investigate the mechanical control of speed in steady undulatory swimming. The roles of body flexural stiffness, driving frequency and driving amplitude were examined; these variables were chosen because of their importance in vibration theory and their hypothesized functions in undulatory swimming. Using a mold of a pumpkinseed sunfish Lepomis gibbosus, we cast three-dimensional vinyl models of four different flexural stiffnesses. We swam the models in a flow tank and powered them via the input of an oscillating sinusoidal bending couple in the horizontal plane at the posterior margin of the neurocranium. To simulate the hydrodynamic conditions of steady swimming, drag and thrust acting on the model were balanced by adjusting flow speed. Under these conditions, the actuated models generated traveling waves of bending. At steady speeds, the motions of the ventral and lateral surfaces of the model were video-taped and analyzed to yield the following response variables: tail-beat amplitude, propulsive wavelength, wave speed and depth of the trailing edge of the caudal fin. Experimental results showed that changes in body flexural stiffness can control propulsive wavelength, wave speed, Froude efficiency and, in consequence, swimming speed. Driving frequency can control tail-beat amplitude, propulsive wavelength, Froude efficiency, relative rate of working and, in consequence, swimming speed. Although there is no significant correlation between rostral amplitude and swimming speed, rostral amplitude can control swimming speed indirectly by controlling tail-beat amplitude and relative power. Compared with live sunfish using undulatory waves at the same speed, models have a lower Froude efficiency. On the basis of the mechanical control of swimming speed in model sunfish, we predict that, in order to swim at fast speeds, live sunfish increase the flexural stiffness of their bodies by a factor of two relative to their passive body stiffness.

Swimming kinematics of fast starts are altered by temperature acclimation in the marine fish Myoxocephalus scorpius

The swimming kinematics of prey capture was studied in short-horned sculpin (Myoxocephalus scorpius L.) acclimated for 6­8 weeks to either 5 °C or 15 °C (12 h:12 h light:dark) using 15 °C-acclimated shrimps as prey. Fish acclimated to 5 °C remained interested in feeding following an acute rise in temperature to 15 °C over 12 h. Prey capture was a stereotyped behaviour consisting of stalking and stationary phases, followed by an S-shaped fast-start (stage 1), a propulsive stroke (stage 2) and a glide of variable duration during which the mouth was expanded and protruded to suck in the prey (stage 3). The duration of the preparatory stroke (half tail-beat, stage 1) was significantly shorter at 15 °C (48.8 ms) than at 5 °C (108.3 ms) in the 5 °C-acclimated sculpin (Q10=2.2). For 5 °C-acclimated fish, at 5 °C, the maximum values for acceleration and velocity along the path travelled by the fish were 16.2 m s-2 and 0.8 m s-1 respectively. Both variables were relatively independent of acute temperature change (Q10=1.1­1.2; P>0.1). At 15 °C, the maximum velocity was 33 % higher and the tail-beat duration of the propulsive stroke was 37 % shorter in 15 °C-acclimated than in 5 °C-acclimated fish. Both stride length and tail-beat amplitude were significantly higher (28 and 23 % respectively) in 15 °C- compared with 5 °C-acclimated sculpin at 15 °C. The results demonstrate plasticity in the major kinematic variables of fast-starts following warm acclimation sufficient to increase the percentage of successful attacks during prey capture from 23.2 to 73.4 %.

UNDULATORY SWIMMING: HOW TRAVELING WAVES ARE PRODUCED AND MODULATED IN SUNFISH (LEPOMIS GIBBOSUS)

We have developed an experimental procedure in which the in situ locomotor muscles of dead fishes can be electrically stimulated to generate swimming motions. This procedure gives the experimenter control of muscle activation and the mechanical properties of the body. Using pumpkinseed sunfish, Lepomis gibbosus, we investigated the mechanics of undulatory swimming by comparing the swimming kinematics of live sunfish with the kinematics of dead sunfish made to swim using electrical stimulation. In electrically stimulated sunfish, undulatory waves can be produced by alternating left­right contractions of either all the axial muscle or just the precaudal axial muscle. As judged by changes in swimming speed, most of the locomotor power is generated precaudally and transmitted to the caudal fin by way of the skin and axial skeleton. The form of the traveling undulatory wave ­ as measured by tail-beat amplitude, propulsive wavelength and maximal caudal curvature ­ can be modulated by experimental control of the body's passive stiffness, which is a property of the skin, connective tissue and axial skeleton.