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The influence of temperature on power production during swimming. I. In vivo length change and stimulation pattern.

Ectothermal animals are able to locomote effectively over a wide range of temperatures despite low temperature reducing the power output of their muscles. It has been suggested that animals recruit more muscle fibres and faster fibre types to compensate for the reduced power output at low temperature, but it is not known how much low temperature actually reduces power output in vivo. 'Optimized' work-loop measurements, which are thought to approximate muscle function in vivo, give a Q(10) of approximately 2.3 for power output of scup (Stenotomus chrysops) red muscle between 10 degrees C and 20 degrees C. However, because of the slower muscle relaxation rate at low temperatures, 'optimizing' work loops requires stimulation duration to be reduced and oscillation frequency to be decreased to obtain maximal power output. Previous fish swimming experiments suggest that similar optimization may not occur in vivo, and this may have substantial consequences in terms of muscle power generation and swimming at low temperatures. To assess more precisely the effects of temperature on muscle performance and swimming, in the present study, we measured the length change, stimulation duration and stimulus phase of red muscle at various positions along scup swimming at several speeds at 10 degrees C and 20 degrees C. In a companion study, we determined the effects of temperature on in vivo power generation by driving muscle fibre bundles through these in vivo length changes and stimulation conditions, and measuring the resulting power output. The most significant finding from the present study is that, despite large differences in the in vivo parameters along the length of the fish (a decrease in stimulus duration, an increase in strain and a negative shift in phase) moving posteriorly, these parameters do not change with temperature. Thus, although the nervous system of fish could, in theory, compensate for slow muscle relaxation by greatly reducing muscle stimulation duration at low temperatures, it does not. This lack of compensation to low temperatures might reflect a potential limitation in neural control.

Animals↗

Swimming kinematics of juvenile kawakawa tuna (Euthynnus affinis) and chub mackerel (Scomber japonicus).

The swimming kinematics of two active pelagic fishes from the family Scombridae were compared to test the hypothesis that the kawakawa tuna (Euthynnus affinis) uses the thunniform mode of locomotion, in which the body is held more rigid and undergoes less lateral movement in comparison with the chub mackerel (Scomber japonicus), which uses the carangiform swimming mode. This study, the first quantitative kinematic comparison of size-matched scombrids, confirmed significantly different swimming kinematics in the two species. Ten kawakawa (15.1-25.5 cm fork length, FL) and eight chub mackerel (14.0-23.4 cm FL), all juveniles, were videotaped at 120 Hz while swimming at several speeds up to their maximum sustained speed at 24 degrees C. Computerized motion analysis was used to digitize specific points on the body in sequential video frames, and kinematic variables were quantified from the progression of the points over time. At a given speed, kawakawa displayed a significantly greater tailbeat frequency, but lower stride length, tailbeat amplitude and propulsive wavelength, than chub mackerel when size effects were accounted for. Midline curvatures subdivided on the basis of X-rays into individual vertebral elements were used to quantify axial bending in a subset of the fish studied. Maximum intervertebral lateral displacement and intervertebral flexion angles were significantly lower along most of the body in kawakawa than in chub mackerel, indicating that the kawakawa undergoes less axial flexion than does the chub mackerel, resulting in lower tailbeat amplitudes. However, lateral movement at the tip of the snout, or yaw, did not differ significantly interspecifically. Despite these differences, the net cost of transport was the same in the two species, and the total cost was higher in the kawakawa, indicating that the tuna juveniles are not more efficient swimmers.

Animals↗

The distribution of NADPH-diaphorase-labelled interneurons and the role of nitric oxide in the swimming system of Xenopus laevis larvae.

The possible involvement of the free radical gas nitric oxide (NO) in the modulation of spinal rhythm-generating networks has been studied using Xenopus laevis larvae. Using NADPH-diaphorase histochemistry, three putative populations of nitric oxide synthase (NOS)-containing cells were identified in the brainstem. The position and morphology of the largest and most caudal population suggested that a proportion of these neurons is reticulospinal. The possible contribution of nitrergic neurons to the control of swimming activity was examined by manipulating exogenous and endogenous NO concentrations in vivo with an NO donor (SNAP, 100-500 micromol l(-)(1)) and NOS inhibitors (l-NAME and l-NNA, 0.5-5 mmol l(-)(1)), respectively. In the presence of SNAP, swim episode duration decreased and cycle period increased, whereas the NOS inhibitors had the opposite effects. We conclude from these data that the endogenous release of NO from brainstem neurons extrinsic to the spinal cord of Xenopus laevis larvae exerts a continuous modulatory influence on swimming activity, functioning like a 'brake'. Although the exact level at which NO impinges upon the swimming rhythm generator has yet to be determined, the predominantly inhibitory effect of NO suggests that the underlying mechanisms of NO action could involve modulation of synaptic transmission and/or direct effects on neuronal membrane properties.

Animals↗

Sustained swimming at low velocity following a bout of exhaustive exercise enhances metabolic recovery in rainbow trout.

Sustained swimming at 0.9 BL s(-)(1), where BL is fork body length, following a bout of exhaustive exercise enhanced recovery of metabolite and acid-base status in rainbow trout compared with fish held in still water. The most striking effect of an active recovery was a total absence of the elevation cortisol concentration typically associated with exhaustive exercise. In fish swimming at 0. 9 BL s(-)(1), plasma cortisol levels averaged 20-25 ng ml(-)(1) throughout the 6 h recovery period. In contrast, plasma cortisol increased to a peak level of 128.4+/-11.2 ng ml(-)(1) (mean +/- s.e. m., N=6) in fish recovering in still water. Muscle glycogen was completely resynthesized and lactate cleared within 2 h of exercise in swimming fish compared with more than 6 h required in the fish held in still water. Similarly, blood lactate level and acid-base status were restored more quickly in the swimming fish. These observations suggest that the prolonged recovery usually associated with exhaustive exercise in rainbow trout is due to elevations in plasma cortisol concentration and that the stimulus for cortisol release is not exercise per se, but rather post-exercise inactivity.

Acid-Base Equilibrium↗

The boundary layer of swimming fish.

Tangential and normal velocity profiles of the boundary layer surrounding live swimming fish were determined by digital particle tracking velocimetry, DPTV. Two species were examined: the scup Stenotomus chrysops, a carangiform swimmer, and the smooth dogfish Mustelus canis, an anguilliform swimmer. Measurements were taken at several locations over the surfaces of the fish and throughout complete undulatory cycles of their propulsive motions. The Reynolds number based on length, Re, ranged from 3x10(3) to 3x10(5). In general, boundary layer profiles were found to match known laminar and turbulent profiles including those of Blasius, Falkner and Skan and the law of the wall. In still water, boundary layer profile shape always suggested laminar flow. In flowing water, boundary layer profile shape suggested laminar flow at lower Reynolds numbers and turbulent flow at the highest Reynolds numbers. In some cases, oscillation between laminar and turbulent profile shapes with body phase was observed. Local friction coefficients, boundary layer thickness and fluid velocities at the edge of the boundary layer were suggestive of local oscillatory and mean streamwise acceleration of the boundary layer. The behavior of these variables differed significantly in the boundary layer over a rigid fish. Total skin friction was determined. Swimming fish were found to experience greater friction drag than the same fish stretched straight in the flow. Nevertheless, the power necessary to overcome friction drag was determined to be within previous experimentally measured power outputs. No separation of the boundary layer was observed around swimming fish, suggesting negligible form drag. Inflected boundary layers, suggestive of incipient separation, were observed sporadically, but appeared to be stabilized at later phases of the undulatory cycle. These phenomena may be evidence of hydrodynamic sensing and response towards the optimization of swimming performance.

Animals↗

The vortex wake of the free-swimming larva and pupa of Culex pipiens (Diptera).

The kinematics and hydrodynamics of free-swimming pupal and larval (final-instar) culicids were investigated using videography and a simple wake-visualisation technique (dyes). In both cases, swimming is based on a technique of high-amplitude, side-to-side (larva) or up-and-down (pupa) bending of the body. The pupa possesses a pair of plate-like abdominal paddles; the larval abdominal paddle consists of a fan of closely spaced bristles which, at the Reynolds numbers involved, behaves like a continuous surface. Wake visualisation showed that each half-stroke of the swimming cycle produces a discrete ring vortex that is convected away from the body. Consecutive vortices are produced first to one side then to the other of the mean swimming path, the convection axis being inclined at approximately 25 degrees away from dead aft. Pupal and larval culicids therefore resemble fish in using the momentum injected into the water to generate thrust. Preliminary calculations for the pupa suggest that each vortex contains sufficient momentum to account for that added to the body with each half-stroke. The possibility is discussed that the side-to-side flexural technique may allow an interaction between body and tail flows in the production of vorticity.

Animals↗

Effects of high intensity exercise training on cardiovascular function, oxygen uptake, internal oxygen transport and osmotic balance in chinook salmon (Oncorhynchus tshawytscha) during critical speed swimming.

To examine cardiorespiratory plasticity, cardiovascular function, oxygen consumption, oxygen delivery and osmotic balance were measured at velocities up to critical swimming speed (Ucrit) in seawater-adapted chinook salmon. We used two groups of fish. The control group had swum continuously for 4 months at a low intensity (0.5 BLs(-1)) and the other was given a high-intensity training regimen (a Ucrit swim test on alternate days) over the same period of time. Compared with available data for other salmonid species, the control group had a higher maximum oxygen consumption (MO2max; 244 micromol O2 min(-1) kg(-1)), cardiac output (Qmax; 65 ml min(-1) kg(-1)) and blood oxygen content (CaO2; 15 ml O2 dl(-1)). Exercise training caused a 50% increase in MO2max without changing either Ucrit or CaO2, even though there were small but significant increases in hematocrit, hemoglobin concentration and relative ventricular mass. During swimming, however, exercise-trained fish experienced a smaller decrease in body mass and muscle moisture, a smaller increase in plasma osmolality, and reduced venous oxygen stores compared with control fish. Consequently, exercise training apparently diminished the osmo-respiratory compromise, but improved oxygen extraction at the tissues. We conclude that the training-induced increase in MO2max provided benefits to systems other than the locomotory system, such as osmoregulation, enabling trained fish to better multitask physiological functions while swimming. Furthermore, because a good interspecific correlation exists between MO2max and arterial oxygen supply (TO2max; r2=0.99) among temperate fish species, it is likely that CaO2 and Qmax are principal loci for cardiorespiratory evolutionary adaptation but not for intraspecific cardiorepiratory plasticity as revealed by high intensity exercise training.

Animals↗

Comparative kinematics of the forelimb during swimming in red-eared slider (Trachemys scripta) and spiny softshell (Apalone spinifera) turtles.

Softshell turtles (Family Trionychidae) possess extensive webbing between the digits of the manus, suggesting that the forelimb may serve as an effective thrust generator during aquatic locomotion. However, the hindlimb has previously been viewed as the dominant propulsive organ in swimming freshwater turtles. To evaluate the potential role of the forelimb in thrust production during swimming in freshwater turtles, we compared the forelimb morphology and three-dimensional forelimb kinematics of a highly aquatic trionychid turtle, the spiny softshell Apalone spinifera, and a morphologically generalized emydid turtle, the red-eared slider Trachemys scripta. Spiny softshells possess nearly twice as much forelimb surface area as sliders for generating drag-based thrust. In addition, although both species use drag-based propulsion, several aspects of forelimb kinematics differ significantly between these species. During the thrust phase of the forelimb cycle, spiny softshells hold the elbow and wrist joints significantly straighter than sliders, thereby further increasing the surface area of the limb that can move water posteriorly and increasing the velocity of the distal portion of the forelimb. These aspects of swimming kinematics in softshells should increase forelimb thrust production and suggest that the forelimbs make more substantial contributions to forward thrust in softshell turtles than in sliders. Spiny softshells also restrict forelimb movements to a much narrower dorsoventral and anteroposterior range than sliders throughout the stroke, thereby helping to minimize limb movements potentially extraneous to forward thrust production. These comparisons demonstrate considerable diversity in the forelimb kinematics of turtles that swim using rowing motions of the limbs and suggest that the evolution of turtle forelimb mechanics produced a variety of contrasting solutions for aquatic specialization.

Animals↗

Function of the heterocercal tail in sharks: quantitative wake dynamics during steady horizontal swimming and vertical maneuvering.

The function of the heterocercal tail in sharks has long been debated in the literature. Previous kinematic data have supported the classical theory which proposes that the beating of the heterocercal caudal fin during steady horizontal locomotion pushes posteroventrally on the water, generating a reactive force directed anterodorsally and causing rotation around the center of mass. An alternative model suggests that the heterocercal shark tail functions to direct reaction forces through the center of mass. In this paper, we quantify the function of the tail in two species of shark and compare shark tail function with previous hydrodynamic data on the heterocercal tail of sturgeon Acipenser transmontanus. To address the two models of shark heterocercal tail function, we applied the technique of digital particle image velocimetry (DPIV) to quantify the wake of two species of shark swimming in a flow tank. Both steady horizontal locomotion and vertical maneuvering were analyzed. We used DPIV with both horizontal and vertical light sheet orientations to quantify patterns of wake velocity and vorticity behind the heterocercal tail of leopard sharks (Triakis semifasciata) and bamboo sharks (Chiloscyllium punctatum) swimming at 1.0Ls(-1), where L is total body length. Two synchronized high-speed video cameras allowed simultaneous measurement of shark body position and wake structure. We measured the orientation of tail vortices shed into the wake and the orientation of the central jet through the core of these vortices relative to body orientation. Analysis of flow geometry indicates that the tail of both leopard and bamboo shark generates strongly tilted vortex rings with a mean jet angle of approximately 30 degrees below horizontal during steady horizontal swimming. The corresponding angle of the reaction force is much greater than body angle (mean 11 degrees ) and the angle of the path of motion of the center of mass (mean approximately 0 degrees ), thus strongly supporting the classical model of heterocercal tail function for steady horizontal locomotion. Vortex jet angle varies significantly with body angle changes during vertical maneuvering, but sharks show no evidence of active reorientation of jet angle relative to body angle, as was seen in a previous study on the function of sturgeon tail. Vortex jet orientation is significantly more inclined than the relatively horizontal jet generated by sturgeon tail vortex rings, demonstrating substantial differences in function in the heterocercal tails of sharks and sturgeon. We present a summary of forces on a swimming shark integrating data obtained here on the tail with previous data on pectoral fin and body function. Body orientation plays a critical role in the overall force balance and compensates for torques generated by the tail. The pectoral fins do not generate lift during steady horizontal locomotion, but play an important hydrodynamic role during vertical maneuvering.

Animals↗

Neuronal control of swimming in the medicinal leech. V. Connexions between the oscillatory interneurones and the motor neurones.

A network of intra- and intersegmental synaptic connexions has been identified in the ventral nerve cord of the leech that links the set of oscillatory interneurones of the central swim oscillator to the motor neurones commanding the swimming rhythm. Excitatory connexions lead from oscillatory interneurones to both excitatory and inhibitory motor neurones, whereas inhibitory connexions lead from oscillatory interneurones to only the inhibitory motor neurones. Connexions leading from a motor neurone back to the oscillatory interneurones were found in only one exceptional case, an inhibitory motor neurone previously known to have access to the central swim oscillator. This network of identified connexions can account reasonably well for the mechanism by which the oscillatory interneurones drive their follower motor neurones into the phasic activity pattern characteristic of the swimming movement.

Action Potentials↗

Neuroethological studies of freely swimming Aplysia brasiliana.

Recently developed neurophysiological and behavioural techniques were used to study swimming in the marine gastropod Aplysia brasiliana. Aplysia swim by bilateral parapodial flapping with an anterior to posterior metachronal wave. Parapodial oscillations are measured from video records. Population recordings from nerves innervating the parapodia during normal swimming in intact Aplysia reveal synchronous phasic activity in large efferent units associated with parapodial opening. Isolated brain studies and stimulation of central pathways in intact animals suggest a central pattern generator. We conclude that the output of the neuronal oscillator that controls parapodial flapping radiates synchronously from each pedal ganglion. The putative command to swim originates within the cerebral ganglia.

Animals↗

Drafting distance in swimming.

PURPOSE: This study investigates the effect of the distance separating the lead and draft swimmers on the metabolic and hydrodynamic responses of the draft swimmer. METHODS: A nondrafting swim of 4 min at 95% of the best 1500-m pace for 11 swimmers was compared with swimming in a drafting position at four different distances directly behind another swimmer (0, 50, 100, and 150 cm). Swimming performance was assessed by stroke rate and stroke length; the metabolic response by oxygen uptake, heart rate, and blood lactate; and the rating of perceived exertion by the Borg scale. Passive drag was assessed at these drafting distances by passive towing. Then, passive drag was measured in six swimmers towed in six lateral drafting positions, with swimmers separated by approximately 40 cm, and then measured in two positions at the rear of the lead swimmer with a reduced lateral distance between swimmers of 50 and 0 cm. RESULTS: Oxygen uptake, heart rate, blood lactate, rating of perceived exertion, and stroke rate were significantly reduced and stroke length was significantly increased in all drafting positions compared with the nondrafting position. For drag, the most advantageous drafting distances were 0 and 50 cm back from the toes of the lead swimmer. Drag was reduced by 21% and 20%, respectively. In lateral drafting, drag was significantly reduced by 6% and 7%, respectively, at 50 and 100 cm back from the hands of the lead swimmer. CONCLUSIONS: Swimming behind another swimmer at a distance between 0 and 50 cm back from the toes was the most advantageous, whereas in lateral drafting the optimal distance was 50-100 cm back from the hands of the lead swimmer.

Adult↗

Protocol for measuring the endurance capacity of mice in an adjustable-current swimming pool.

We re-examined the methods used in an adjustable-current swimming pool for evaluating the endurance capacity of mice to improve the sensitivity and reproducibility. We found that the BALB/c strain was most suitable to minimize the variation in time taken to swim to the point of fatigue. We found that precise adjustment of the apparatus and the use of three primary swimming trials to select mice before a study made the swimming time more uniform. This procedure was used to establish a more precise evaluation protocol.

Animals↗

Capsiate, a nonpungent capsaicin analog, increases endurance swimming capacity of mice by stimulation of vanilloid receptors.

We investigated the effect of capsiate, a nonpungent natural capsaicin analog, on the swimming capacity of mice in an adjustable-current water pool. Male BALB/c mice orally given capsiate (10 mg/kg) were able to keep swimming longer before exhaustion than the control mice. After 30 min of swimming, the residual glycogen in the gastrocnemius muscle was higher, the serum free fatty acid concentration tended to be higher, and the serum lactic acid concentration was significantly lower in the capsiate-administered mice. The value for the respiratory exchange ratio of the capsiate group was significantly lower during both resting and treadmill running. These physiological differences were abolished by administering the vanilloid receptor antagonist, capsazepin (0.17 mmol/kg, i.p.). The mice were not averse to the capsiate solution during a 4-h two-bottle choice test. These results suggest that the oral administration of capsiate enhanced fat oxidation and spared carbohydrate utilization, and consequently increased the endurance swimming capacity of the mice via stimulation of their vanilloid receptors. Practical application of capsiate is expected.

Animals↗

Rapidly measured indicators of recreational water quality are predictive of swimming-associated gastrointestinal illness.

Standard methods to measure recreational water quality require at least 24 hr to obtain results, making it impossible to assess the quality of water within a single day. Methods to measure recreational water quality in <or=2 hr have been developed. Application of rapid methods could give considerably more accurate and timely assessments of recreational water quality. We conducted a prospective study of beachgoers at two Great Lakes beaches to examine the association between recreational water quality, obtained using rapid methods, and gastrointestinal (GI) illness after swimming. Beachgoers were asked about swimming and other beach activities and 10-12 days later were asked about the occurrence of GI symptoms. We tested water samples for Enterococcus and Bacteroides species using the quantitative polymerase chain reaction (PCR) method. We observed significant trends between increased GI illness and Enterococcus at the Lake Michigan beach and a positive trend for Enterococcus at the Lake Erie beach. The association remained significant for Enterococcus when the two beaches were combined. We observed a positive trend for Bacteroides at the Lake Erie beach, but no trend was observed at the Lake Michigan beach. Enterococcus samples collected at 0800 hr were predictive of GI illness that day. The association between Enterococcus and illness strengthened as time spent swimming in the water increased. This is the first study to show that water quality measured by rapid methods can predict swimming-associated health effects.

Adult↗

Changes in skeletal muscle composition in response to swimming training for young horses.

To investigate muscular adaptation to swimming training in young horses, 18 two-year-old Thoroughbred horses were trained in a program which included both running and swimming, and the changes in skeletal muscle composition during the training period were evaluated histochemically. The horses were divided into the following three groups; Group A, trained by running only: Group B, trained by running plus a gradual increase in swimming; Group C, trained by running plus constant swimming. In Groups B and C, fast twitch-low oxidative (FT) fibers tended to decrease, while fast twitch-high oxidative (FTH) fibers to increase with the training progressed. The tendency was more remarkable in Group B. Therefore, in these two groups, it was suggested that the horses' aerobic capacity at the muscular level was improved significantly.

Aging↗

Characteristics of respiratory function during swimming exercise in thoroughbreds.

Equine respiratory patterns during swimming were examined in five normal horses. The experiment included a preliminary warming-up stage and 6 circuits of swimming around an annular pool of a 50-meter-circumference. The horses were examined for respiratory rates, intratracheal pressures, inspiratory time (TI), expiratory time (TE), respiratory cycle (T; TI + TE), heart rates, blood lactate concentrations, hematocrit and blood gases. The respiratory rates were maintained around 25/min. Blood gas values changed significantly during swimming. The intratracheal pressures during expiration and inspiration increased significantly with exercise duration compared to the immediately after the warming-up stage. The duty ratio (TI/T) averaged 0.33, which implied that the expiratory time was roughly doubled the inspiratory time. We considered that a longer expiratory time may limit sudden collapse of airways by water pressure during swimming and prevent a radical decrease of air space volume, thus maintains buoyancy.

Animals↗

A trial to detect behavioral changes by swimming stress.

We investigated behavioral changes in mice in a light/dark test after forced swimming stress in a streaming pool. Locomotion and rearing behavior in mice in a light/dark test was suppressed immediately after the swimming and gradually regained. Shuttle crossing between light and dark zones was also suppressed but time spent in a light zone was not. The suppression of locomotor activity by the swimming was not observed 20 min after it. Naloxone (10 mg/kg, i.p.) suppressed recovery of locomotor activity after the swimming while it did not affect a light/dark test in nonswimming mice. These results suggest that this model is useful for the study of behavioral and psychological changes after exercise, and that opioidergic systems may be involved in regaining spontaneous locomotor activity.

Animals↗