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Specificity of swim training on maximum oxygen uptake.

The present study was designed to evaluate the specificity-generality of the cardiorespiratory adaptation to swim training. Fifteen male, college age, recreational swimmers utilized interval swim training procedures 1 h/day, 3 days/wk for 10 wk. Maximum physiological measures (Vo2, Ve, HR, R, and work time) were determined prior to and following swim training during treadmill running and tethered swimming Vo2max tests. Identical measures were made on 15 control subjects who did not participate in any form of training. As a result of training, the experimental subjects significantly increased (all P is less than 0.01) their swimming Vo2max (380 ml/min) max Ve (14.9 l/min,btps) and max swim time (4.0 min), and significantly decreased (P is less than 0.05) their max HR (3.5 beats/min). However, there was no significant improvement in Vo2max when the same subjects were evaluated by the treadmill running test. Differences in Vo2max and associated measures during running and swimming tests remained essentially unchanged for control subjects. The results of the present study clearly demonstrate the specificity of the cardiorespiratory adaptation to swim training in male recreational swimmers.

Adaptation, Physiological↗

Surfactant homeostasis in the rat lung during swimming exercise.

Swimming rats for up to 2 h in water at 34 +/- 1 degree C increased the rate of breathing by 60% and tidal volume by 200-300%. In each case we infused 20 muCi X kg-1 [methyl-3H]choline chloride, via a caudal vein, 3 h prior to the end of swimming. Maximum specific activity of tissue phospholipid (PL) and alveolar PL (PLalv) occurred in 1 and 12 h, respectively. Total PLalv, specific activity of PLalv and the percentage of total PL released (%A/T) increased within 10 min of start of swimming and were sustained for at least 2 h of swimming [PLalv in mg X g dry lung-1: 5 s swim, 7.3 +/- 0.96 (mean +/- SD; n = 15 rats); 1-h swim, 10.1 +/- 1.1 (n = 23 rats)]. After a 1-h swim, PLalv returned to control within 4 h. Pretreatment with propranolol hydrochloride (10 mg X kg-1) (P), atropine methyl nitrate (3 mg X kg-1) (A), indomethacin (15 mg X kg-1), and cyproheptadine (1 mg X kg-1) did not alter the increase in PLalv with swimming, however, both P and A reduced the increase in specific activity of PLalv and %A/T. We suggest that the exercise releases surfactant from two pools: a readily released pool that responds to direct distortion of the alveolar type II cell and a pool that is under sympathetic nervous control.

Animals↗

Activities of identified interneurons, motoneurons, and muscle fibers during fictive swimming in the lamprey and effects of reticulospinal and dorsal cell stimulation.

1. Application of D-glutamate to the isolated spinal cord of the lamprey produces phasic activity in ventral roots, which is similar to that of the muscles of the intact swimming animal (5,18). Therefore, the isolated spinal cord may be used as a convenient model for the investigation of the generation of locomotor rhythms in a vertebrate. 2. Almost all slow muscle fibers exhibited excitatory junctional potentials (EJPs) during swimming activity. The number of EJPs per cycle increased with the intensity of ventral root (VR) bursting. Few twitch fibers were active, and these fired action potentials only during high intensities of VR bursts. 3. As was found by Russell and Wallén (25), myotomal motoneurons had oscillating membrane potentials during fictive swimming which, on the average, reached a peak depolarization in the middle of the VR burst (phi = 0.21 +/- 0.05; phi = 0 is defined as the onset of the VR burst, and the duration of the cycle is set equal to 1). Membrane potential oscillations in fin motoneurons were antiphasic to those of nearby myotomal motoneurons (peak depolarization phi = 0.68 +/- 0.05). 4. Lateral interneurons had oscillating membrane potentials in synchrony with those of myotomal motoneurons (peak depolarization phi = 0.21 +/- 0.10). Interneurons with axons projecting contralaterally and caudally (CC interneurons) had oscillating membrane potentials that peaked significantly earlier in the cycle (peak depolarization phi = 0.06 +/- 0.12). 5. Edge cells were only weakly modulated during fictive swimming. Their peak depolarizations occurred near the end of the VR burst (phi = 0.33 +/- 0.10). Most giant interneurons were not phasically modulated during fictive swimming. 6. Repetitive intracellular stimulation of Müller cells during fictive swimming generally evoked an increased burst intensity in ipsilateral VRs and a decreased burst intensity in contralateral VRs. The cells M3, B1, and B2 also produced increases or decreases in the frequency of VR bursts. Repetitive intracellular stimulation of sensory dorsal cells could also change the intensities and timing of VR bursts. 7. This study is an initial survey of lamprey spinal interneurons that participate in swimming activity. Lateral interneurons and CC interneurons are active during fictive swimming and probably help coordinate the undulations of the body, but their roles in pattern generation are not known. The central pattern generator is subject to modification by descending and sensory inputs.

Action Potentials↗

Swimming pattern in intact and decerebrated stingrays.

Stingrays swim with a rostral-to-caudal wave of elevation and depression of the enlarged pectoral fins. The present study compares the timing relations of swimming before and after decerebration in the same animals using electromyograms (EMGs). EMGs were recorded from several freely swimming, intact animals. Most recordings, however, were from stingrays suspended by vertebral clamps. Activity recorded from restrained animals exhibited the same timing relations as freely swimming animals. Therefore, the restrained preparation was used to compare locomotor rhythm in intact and decerebrate stingrays. The delay from the onset of activity recorded at a rostral segment to the onset of activity at successively caudal recording sites (intersegmental delay) was linearly related to the extent of the segmental separation. This relation was not changed by decerebration. Both the intersegmental delay and the EMG burst duration were linearly related to the period of the swim cycle. The intercepts for both relations fell around the origin. Therefore, stingray swimming demonstrates constant phase coupling. Neither the slopes nor the intercepts of the coupling relations were altered by decerebration. Histological examination of the fins indicates that at least two types of muscles exist. Superficial muscle is activated during slow swimming, while the deeper muscle is recruited during faster swimming.

Animals↗

Mechanisms of pattern generation underlying swimming in Tritonia. IV. Gating of central pattern generator.

Swimming behavior in the marine mollusc Tritonia diomedea is episodic, consisting of a series of alternating dorsal and ventral flexions initiated by a brief sensory stimulus. The swim motor pattern is generated by a network formed of four groups of premotor interneurons: cerebral cell 2 (C2), dorsal swim interneurons (DSIs), and two types of ventral swim interneurons (VSI-A and VSI-B). The initiation and maintenance of swimming depends on the establishment of a long-lasting ramp depolarization in both the premotor, pattern-generating interneurons, and the motor neurons (i.e., flexion neurons). Voltage clamp was used to measure the membrane current responsible for the ramp depolarization. In all cell classes the current had two components: a tonic inward current, which decayed as the swim progressed, and phasic inward current waves, which provided the synaptic drive during each swim burst. The ramp current in the flexion neurons and in C2 was generated largely by activity within the interneuronal pattern-generating network (PGN). The ramp current could be mimicked by driving activity in the pattern-generating interneurons. In VSI-B, the tonic component of the ramp current was independent of activity within the PGN and appeared to be derived from the long-lasting effect of an extrinsic input. The phasic components of the ramp, however, were dependent on PGN activity. The phasic inward current waves were blocked when pattern generation was prevented. In addition, phasic inward currents similar to those occurring during swimming could be produced by driving the C2. The tonic component of the ramp current in a DSI was dependent both on extrinsic inputs and PGN activity. Extrinsic inputs appeared to control the first 10-15 s of the tonic current. At longer times, activity within the DSI population itself maintained the ramp current. When one DSI was driven in a quiescent preparation, all other DSIs were inhibited, yet the DSIs are known to be coupled by monosynaptic, reciprocal excitatory synapses. This effect could be explained by the action of an unidentified inhibitory interneuron (I-neuron), which was excited by DSIs and in turn inhibited all other DSIs. The DSIs were therefore coupled reciprocally by both monosynaptic excitation and polysynaptic inhibition. Activity in C2 switched the DSI-DSI interaction from inhibition to excitation by inhibiting the I-neuron.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

Synaptic potentials in motoneurons during fictive swimming in spinal Xenopus embryos.

Embryos spinalized at the 3rd to 6th postotic myotome and immobilized in 10(-4) M tubocurarine can respond to a brief skin stimulus with motor root activity suitable for swimming. Embryos spinalized at the more caudal levels give shorter episodes of fictive swimming. We have previously described the synaptic inputs to motoneurons during fictive swimming in intact embryos (23). In the present paper we look to see if similar synaptic inputs are present in spinal embryos and are therefore spinal in origin. All motoneuron firing during fictive swimming is associated with a tonic depolarization that falls away slowly once firing stops, is increased by hyperpolarizing current, and is reduced by depolarizing current. A slow depolarizing potential evoked by lower levels of skin stimulation has similar properties and rate of fall. In 1-2 mM PDA, an excitatory amino acid antagonist, only a small remnant of the depolarization remains, and motoneuron firing stops. The NMDA antagonist 50 microM APV reduces the depolarization less but also blocks firing. Motoneurons fire one spike per swimming cycle, in phase with nearby motor root discharge. Spikes are preceded by a depolarizing prepotential. This increases with hyperpolarizing current, which can block the spike to reveal an underlying depolarizing potential. In phase with motor root discharge on the opposite side of the body, motoneurons receive a midcycle inhibitory postsynaptic potential, which increases with depolarizing current, decreases with hyperpolarizing current, and is blocked by 10(-6) M strychnine. Strychnine, 5 X 10(-7) M, leads first to broadening of motor root bursts then to loss of the alternating swimming pattern of activity, which is replaced by synchronous bursts on both sides of the body. We conclude that the synaptic inputs to motoneurons during fictive swimming in spinal embryos are very similar in properties and pharmacology to those in intact embryos. These inputs, including the tonic depolarization always associated with motoneuron firing during swimming, must be at least partly spinal in origin.

Amino Acids↗

Struggling and flumazenil effects in the swimming test are related to the level of anxiety in mice.

The possible involvement of anxiety and learning/memory processes in escape-directed (struggling) behavior in a two-trial swimming test was investigated in mice, as well as the differential effects that low doses of flumazenil (a benzodiazepine receptor antagonist) could display depending on the animals' anxiety levels. Mice showing less anxiety in the plus-maze test exhibited less struggling behavior in the first swimming trial than the more anxious animals, suggesting a relationship between anxiety and struggling behavior in the swimming test. Flumazenil (5 mg/kg) given before the first swimming trial displayed differential effects depending upon the animals' anxiety levels. Thus, it increased struggling behavior in the first swimming trial in 'low-anxiety' mice whereas the opposite tendency was observed in 'high-anxiety' animals. Struggling decreased in the second swimming trial in all the animals, giving support to the involvement of learning/memory processes in the two-trial swimming test. That reduction in escape-directed behavior was greater in animals treated with flumazenil before the first swimming session, thus indicating a slight enhancement of retention.

Animals↗

The effect of swimming on patients with ischemic heart disease.

Swimming is frequently recommended for cardiac rehabilitation, but little is known of its physiologic consequences in ischemic heart disease. Eight males who had had a myocardial infarction 8-17 months before the study were exercised to exhaustion or angina with 10 W/min-1 ramp on a cycle ergometer in sitting and supine positions. Oxygen uptake (VO2) was continuously measured to monitor the physiologic power requirement. All eight patients were taking beta blockers and four were taking digoxin. During sitting cycling, angina occurred in four and ST depression in five; during supine cycling, angina occurred in five and ST depression in six. VO2 was then measured while they swam at their own comfortable speed (mean 0.43 m/sec-1) in a swimming flume at water temperatures of 25.5 degrees C and 18 degrees C. In six, the water speed was gradually increased until they were limited by symptoms. Comfortable swimming at 25.5 degrees C was 87% (1.28 1/min-1) and at 18 degrees C 89% (1.30 1/min-1) of sitting peak VO2, while heart rates were 92% and 91% respectively. The mean peak VO2 and heart rate did not differ significantly between bicycle and swim tests (peak VO2 sitting 1.49 +/- 0.23, supine 1.42 +/- 0.24, 25.5 degrees C 1.60 +/- 0.17, 18 degrees C 1.52 +/- 0.19 1/min-1). Only two patients reported angina while swimming in warm water and one in cold water, although ST depression occurred in six in both swims. The subjective comfort and large muscle groups involved make swimming a good exercise, but the high relative energy cost and failure to identify ischemic symptoms indicate caution in cardiac patients, especially if their swimming skills are poor.

Aged↗

Toxic cocaine- and convulsant-induced modification of forced swimming behaviors and their interaction with ethanol: comparison with immobilization stress.

BACKGROUND: Swimming behaviors in the forced swimming test have been reported to be depressed by stressors. Since toxic convulsion-inducing drugs related to dopamine [cocaine (COC)], benzodiazepine [methyl 6,7-dimethoxy-4-ethyl-beta-carboline-carboxylate (DMCM)], gamma-aminobutyric acid (GABA) [bicuculline (BIC)], and glutamate [N-methyl-D-aspartate (NMDA)] receptors can function as stressors, the present study compared their effects on the forced swimming behaviors with the effects of immobilization stress (IM) in rats. Their interactions with ethanol (EtOH), the most frequently coabused drug with COC which also induces convulsions as withdrawal symptoms but interferes with the convulsions caused by other drugs, were also investigated. RESULTS: Similar to the IM (10 min) group, depressed swimming behaviors (attenuated time until immobility and activity counts) were observed in the BIC (5 mg/kg IP) and DMCM (10 mg/kg IP) groups at the 5 h time point, after which no toxic behavioral symptoms were observed. However, they were normalized to the control levels at the 12 h point, with or without EtOH (1.5 g/kg IP). In the COC (60 mg/kg IP) and NMDA (200 mg/kg IP) groups, the depression occurred late (12 h point), and was normalized by the EtOH cotreatment. At the 5 h point, the COC treatment enhanced the swimming behaviors above the control level. CONCLUSIONS: Although the physiological stress (IM), BIC, and DMCM also depressed the swimming behaviors, a delayed occurrence and EtOH-induced recovery of depressed swimming were observed only in the COC and NMDA groups. This might be correlated with the previously-reported delayed responses of DA and NMDA neurons rather than direct effects of the drugs, which could be suppressed by EtOH. Furthermore, the characteristic psychostimulant effects of COC seemed to be correlated with an early enhancement of swimming behaviors.

Animals↗

Effects of postnatal cocaine exposure and environmental enrichment on rat behavior in a forced swim test.

This study examined the effects of environmental enrichment on rats exposed to cocaine during the first month of life, in several categories of behavior observed in a forced swim test. Wistar rats were divided in four groups. The first included pups that were subjected to injections of cocaine hydrochloride (15 mg/kg body weight/day, subcutaneously, in two daily doses, from postnatal days 1 to 27) and reared in an enriched environment (CocEE); the second, pups that were subjected to injections of cocaine (as previously described) and reared in a standard environment (CocSE); the third, pups that were subjected to saline injections and reared in an enriched environment (SalEE); the fourth, pups that were subjected to saline injections and reared in a standard environment (SalSE). On postnatal days 26 and 27, rats were tested in a swimming pool in two 5-min sessions. The categories of behavior studied in this work were: fast swim, slow swim, struggling, diving, and immobility. Results showed that postnatal cocaine exposure decreased the time spent on fast swim during the two sessions and increased the immobility behavior during the second session in CocSE pups compared with SalSE pups. SalEE pups increased the time spent in fast swim, slow swim, and diving, and decreased the time spent in struggling and immobility during the two sessions compared with SalSE pups. CocEE animals spent more time in fast swim and struggling and less the time in immobility compared with CocSE pups. The present results suggest that postnatal cocaine exposure affects the ability of these animals to cope with stressful situations, and that environmental enrichment seems to enable the rats to adopt a more active strategy, one that allows them to better cope with this particular stress situation.

Animals↗

Swimming performance studies on the eastern Pacific bonito Sarda chiliensis, a close relative of the tunas (family Scombridae) I. Energetics.

A large swim tunnel respirometer was used to quantify the swimming energetics of the eastern Pacific bonito Sarda chiliensis (tribe Sardini) (45-50 cm fork length, FL) at speeds between 50 and 120 cm s(-1) and at 18+/-2 degrees C. The bonito rate of oxygen uptake ((O(2)))-speed function is U-shaped with a minimum (O(2)) at 60 cm s(-1), an exponential increase in (O(2)) with increased speed, and an elevated increase in (O(2)) at 50 cm s(-1) where bonito swimming is unstable. The onset of unstable swimming occurs at speeds predicted by calculation of the minimum speed for bonito hydrostatic equilibrium (1.2 FL s(-1)). The optimum swimming speed (U(opt)) for the bonito at 18+/-2 degrees C is approximately 70 cm s(-1) (1.4 FL s(-1)) and the gross cost of transport at U(opt) is 0.27 J N(-1) m(-1). The mean standard metabolic rate (SMR), determined by extrapolating swimming (O(2)) to zero speed, is 107+/-22 mg O(2) kg(-1) h(-1). Plasma lactate determinations at different phases of the experiment showed that capture and handling increased anaerobic metabolism, but plasma lactate concentration returned to pre-experiment levels over the course of the swimming tests. When adjustments are made for differences in temperature, bonito net swimming costs are similar to those of similar-sized yellowfin tuna Thunnus albacares (tribe Thunnini), but the bonito has a significantly lower SMR. Because bonitos are the sister group to tunas, this finding suggests that the elevated SMR of the tunas is an autapomorphic trait of the Thunnini.

Animals↗

Swimming of larval zebrafish: ontogeny of body waves and implications for locomotory development.

Fish larvae, like most adult fish, undulate their bodies to propel themselves. A detailed kinematic study of the larval body wave is a prerequisite to formulate a set of functional requirements that the locomotor system must fulfil to generate the observed swimming kinematics. Lateral displacement and curvature profiles were obtained for zebrafish (Danio rerio) larvae at 2-21 days post-fertilisation for three swimming behaviours (cyclic swimming, slow starts and fast startle responses) using high-speed video. During cyclic swimming, fish larvae maintain tail beat frequencies of up to 100 Hz. The corresponding longitudinal strains, estimated from the peak curvatures of the midline, reach up to 0.19 in superficial tissue. The strain rate can reach 120 s(-1). The wave of curvature travels along the body at a near-constant rate. Posterior to the stiff head, body-length-specific curvature is high and rises gently along the entire trunk to a maximum value of 6. Burst-and-coast swimming generates similar peak curvatures to cyclic swimming, but curvature rises more steeply from head to tail. Fish larvae exhibit phase shifts of 57-63 degrees between the wave of lateral displacement and the wave of curvature, resulting in a 1:1.2 ratio of body wave length to curvature wave length. During C-starts, muscle strain can reach 0.19 and superficial longitudinal strain rates approach 30 s(-1). Fish larvae do not initiate their escape response with a standing wave of curvature, although their C-starts approach a standing wave as the larvae grow older. The performance demands derived from swimming kinematics suggest that larval axial muscles have very short contraction cycles (10 ms), experience considerable strains (up to 0.2) and strain rates (up to 30 s(-1) in white muscle fibres) yet are able to power swimming for several seconds.

Animals↗

Strouhal numbers and optimization of swimming by odontocete cetaceans.

Swimming efficiencies of fish and cetaceans have been related to a certain synchrony between stroke cycle frequency, peak-to-peak tail/fluke amplitude and mean swimming speed. These kinematic parameters form a non-dimensional wake parameter, referred to as a Strouhal number, which for the range between 0.20 and 0.40 has been associated with enhanced swimming efficiency for fish and cetaceans. Yet to date there has been no direct experimental substantiation of what Strouhal numbers are preferred by swimming cetaceans. To address this lack of data, a total of 248 Strouhal numbers were calculated for the captive odontocete cetaceans Tursiops truncatus, Pseudorca crassidens, Orcinus orca, Globicephala melaena, Lagenorhynchus obliquidens and Stenella frontalis. Although the average Strouhal number calculated for each species is within the accepted range, considerable scatter is found in the data both within species and among individuals. A greater proportion of Strouhal values occur between 0.20 and 0.30 (74%) than the 0.25-0.35 (55%) range predicted for maximum swimming efficiency. Within 0.05 Strouhal increments, the greatest number of Strouhal values was found between 0.225 and 0.275 (44%). Where propulsive efficiency data were available (Tursiops truncatus, Pseudorca crassidens, Orcinus orca), peak swimming efficiency corresponded to this same Strouhal range. The odontocete cetacean data show that, besides being generally limited to a range of Strouhal numbers between 0.20 and 0.40, the kinematic parameters comprising the Strouhal number provide additional constraints. Fluke-beat frequency normalized by the ratio of swimming speed to body length was generally restricted from 1 to 2, whereas peak-to-peak fluke amplitude normalized by body length occurred predominantly between 0.15 and 0.25. The results indicate that the kinematics of the propulsive flukes of odontocete cetaceans are not solely dependent on Strouhal number, and the Strouhal number range for odontocete cetaceans occurs at slightly ( approximately 20%) lower values than previously predicted for maximum swimming efficiency.

Animals↗

Propulsive force calculations in swimming frogs. I. A momentum-impulse approach.

Frogs are animals that are capable of locomotion in two physically different media, aquatic and terrestrial. A comparison of the kinematics of swimming frogs in a previous study revealed a difference in propulsive impulse between jumping and swimming. To explore this difference further, we determined the instantaneous forces during propulsion in swimming using an impulse-momentum approach based on DPIV flow data. The force profile obtained was compared with force profiles obtained from drag-thrust equilibrium of the centre of mass and with the force profiles generated during jumping. The new approach to quantifying the instantaneous forces during swimming was tested and proved to be a valid method for determining the external forces on the feet of swimming frogs. On the kinematic profiles of swimming, leg extension precedes propulsion. This means that it is not only the acceleration of water backwards that provides thrust, but also that the deceleration of water flowing towards the frog as a result of recovery accelerates the centre of mass prior to leg extension. The force profile obtained from the impulse-momentum approach exposed an overestimation of drag by 30% in the drag-thrust calculations. This means that the difference in impulse between jumping and swimming in frogs is even larger than previously stated. The difference between the force profiles, apart from a slightly higher peak force during jumping, lies mainly in a difference in shape. During swimming, maximal force is reached early in the extension phase, 20% into it, while during jumping, peak force is attained at 80% of the extension phase. This difference is caused by a difference in inter-limb coordination.

Animals↗

A hydrodynamic analysis of fish swimming speed: wake structure and locomotor force in slow and fast labriform swimmers.

Past study of interspecific variation in the swimming speed of fishes has focused on internal physiological mechanisms that may limit the ability of locomotor muscle to generate power. In this paper, we approach the question of why some fishes are able to swim faster than others from a hydrodynamic perspective, using the technique of digital particle image velocimetry which allows measurement of fluid velocity and estimation of wake momentum and mechanical forces for locomotion. We investigate the structure and strength of the wake in three dimensions to determine how hydrodynamic force varies in two species that differ markedly in maximum swimming speed. Black surfperch (Embiotoca jacksoni) and bluegill sunfish (Lepomis macrochirus) swim at low speeds using their pectoral fins exclusively, and at higher speeds switch to combined pectoral and caudal fin locomotion. E. jacksoni can swim twice as fast as similarly sized L. macrochirus using the pectoral fins alone. The pectoral fin wake of black surfperch at all speeds consists of two distinct vortex rings linked ventrally. As speed increases from 1.0 to 3.0 L s(-)(1), where L is total body length, the vortex ring formed on the fin downstroke reorients to direct force increasingly downstream, parallel to the direction of locomotion. The ratio of laterally to downstream-directed force declines from 0.93 to 0.07 as speed increases. In contrast, the sunfish pectoral fin generates a single vortex ring per fin beat at low swimming speeds and a pair of linked vortex rings (with one ring only partially complete and attached to the body) at maximal labriform speeds. Across a biologically relevant range of swimming speeds, bluegill sunfish generate relatively large lateral forces with the paired fins: the ratio of lateral to downstream force remains at or above 1.0 at all speeds. By increasing wake momentum and by orienting this momentum in a direction more favorable for thrust than for lateral force, black surfperch are able to swim at twice the speed of bluegill sunfish using the pectoral fins. In sunfish, without a reorientation of shed vortices, increases in power output of pectoral fin muscle would have little effect on maximum locomotor speed. We present two hypotheses relating locomotor stability, maneuverability and the structure of the vortex wake. First, at low speeds, the large lateral forces exhibited by both species may be necessary for stability. Second, we propose a potential hydrodynamic trade-off between speed and maneuverability that arises as a geometric consequence of the orientation of vortex rings shed by the pectoral fins. Bluegill sunfish may be more maneuverable because of their ability to generate large mediolateral force asymmetries between the left- and right-side fins.

Animals↗

Swimming mechanics and behavior of the shallow-water brief squid Lolliguncula brevis.

Although squid are among the most versatile swimmers and rely on a unique locomotor system, little is known about the swimming mechanics and behavior of most squid, especially those that swim at low speeds in inshore waters. Shallow-water brief squid Lolliguncula brevis, ranging in size from 1.8 to 8.9 cm in dorsal mantle length (DML), were placed in flumes and videotaped, and the data were analyzed using motion-analysis equipment. Flow visualization and force measurement experiments were also performed in water tunnels. Mean critical swimming speeds (U(crit)) ranged from 15.3 to 22.8 cm s(-1), and mean transition speeds (U(t); the speed above which squid swim exclusively in a tail-first orientation) varied from 9.0 to 15.3 cm s(-1). At low speeds, negatively buoyant brief squid generated lift and/or improved stability by positioning the mantle and arms at high angles of attack, directing high-speed jets downwards (angles >50 degrees ) and using fin activity. To reduce drag at high speeds, the squid decreased angles of attack and swam tail-first. Fin motion, which could not be characterized exclusively as drag- or lift-based propulsion, was used over 50-95 % of the sustained speed range and provided as much as 83.8 % of the vertical and 55.1 % of the horizontal thrust. Small squid (<3.0 cm DML) used different swimming strategies from those of larger squid, possibly to maximize thrust benefits from vortex ring formation. Furthermore, brief squid employed various unsteady behaviors, such as manipulating funnel diameter during jetting, altering arm position and swimming in different orientations, to boost swimming performance. These results demonstrate that locomotion in slow-swimming squid is complex, involving intricate spatial and temporal interactions between the mantle, fins, arms and funnel.

Animals↗

Growth and metabolism of larval zebrafish: effects of swim training.

Larval zebrafish (Danio rerio) of three different age classes ('yolk-sac' larvae, 96 h; 'swim-up' larvae, 9 days old; and 'free-swimming' larvae, 21 days old) were trained for 2, 6 and 11 days, respectively, to swim at 0 body lengths per second (BL s(-1)), 2 BL s(-1) and 5 BL s(-1). Survival was significantly diminished in larvae trained at 5 BL s(-1) compared to controls (0 BL s(-1)). Although training produced no significant differences in mass and length, the youngest larvae absorbed their yolk at a faster rate during training. Routine oxygen consumption ((MO(2)r)) and mass-specific routine oxygen consumption ((MO(2)r,m)) were not significantly affected by chronic training in the yolk-sac larvae and swim-up larvae. However, trained free-swimming larvae had a significantly higher (MO(2)r) (after 11 days of training) and (MO(2)r,m) (after 8 and 11 days of training) compared to control larvae. Trained free-swimming larvae consumed significantly less oxygen during swimming compared to control larvae, as measured by closed-system respirometry. Trained yolk-sac larvae exposed to increasing hypoxia levels were more effective O(2) regulators. Additionally, training enhanced survival during exposure to extreme hypoxia in all age groups. Thus, physiological acclimation to chronic swimming occurs in the earliest stages of life in the zebrafish.

Adaptation, Physiological↗

Effects of longitudinal body position and swimming speed on mechanical power of deep red muscle from skipjack tuna (Katsuwonus pelamis).

The mechanical power output of deep, red muscle from skipjack tuna (Katsuwonus pelamis) was studied to investigate (i) whether this muscle generates maximum power during cruise swimming, (ii) how the differences in strain experienced by red muscle at different axial body locations affect its performance and (iii) how swimming speed affects muscle work and power output. Red muscle was isolated from approximately mid-way through the deep wedge that lies next to the backbone; anterior (0.44 fork lengths, ANT) and posterior (0.70 fork lengths, POST) samples were studied. Work and power were measured at 25 degrees C using the work loop technique. Stimulus phases and durations and muscle strains (+/- 5.5 % in ANT and +/- 8 % in POST locations) experienced during cruise swimming at different speeds were obtained from previous studies and used during work loop recordings. In addition, stimulus conditions that maximized work were determined. The stimulus durations and phases yielding maximum work decreased with increasing cycle frequency (analogous to tail-beat frequency), were the same at both axial locations and were almost identical to those used by the fish during swimming, indicating that the muscle produces near-maximal work under most conditions in swimming fish. While muscle in the posterior region undergoes larger strain and thus produces more mass-specific power than muscle in the anterior region, when the longitudinal distribution of red muscle mass is considered, the anterior muscles appear to contribute approximately 40% more total power. Mechanical work per length cycle was maximal at a cycle frequency of 2-3 Hz, dropping to near zero at 15 Hz and by 20-50% at 1 Hz. Mechanical power was maximal at a cycle frequency of 5 Hz, dropping to near zero at 15 Hz. These fish typically cruise with tail-beat frequencies of 2.8-5.2 Hz, frequencies at which power from cyclic contractions of deep red muscles was 75-100% maximal. At any given frequency over this range, power using stimulation conditions recorded from swimming fish averaged 93.4 +/- 1.65% at ANT locations and 88.6 +/- 2.08% at POST locations (means +/- S.E.M., N=3-6) of the maximum using optimized conditions. When cycle frequency was held constant (4 Hz) and strain amplitude was increased, work and power increased similarly in muscles from both sample sites; work and power increased 2.5-fold when strain was elevated from +/- 2 to +/- 5.5%, but increased by only approximately 12% when strain was raised further from +/- 5.5 to +/- 8%. Taken together, these data suggest that red muscle fibres along the entire body are used in a similar fashion to produce near-maximal mechanical power for propulsion during normal cruise swimming. Modelling suggests that the tail-beat frequency at which power is maximal (5 Hz) is very close to that used at the predicted maximum aerobic swimming speed (5.8 Hz) in these fish.

Animals↗