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Identification and characterization of cerebral ganglion neurons that induce swimming and modulate swim-related pedal ganglion neurons in Aplysia brasiliana.

1. We have identified and characterized a family of several pairs of neurons in the cerebral ganglion of Aplysia brasiliana that are capable of inducing, maintaining, or modulating a motor program that underlies swim locomotion in this marine mollusk. We have operationally defined these cells as command neurons (CNs) for swimming. 2. The command cells occur in bilateral pairs in the cerebral ganglion and make direct and indirect outputs to neurons in the pedal ganglia, including motor neurons, a central pattern generator circuit, and modulatory neurons that enhance muscle contractions during swimming. Several of the CNs are sufficient individually to induce the swim motor program (SMP), all receive sensory feedback from the periphery, and several interconnect with other swim-related CNs. 3. Tonic discharges of approximately 10 Hz in CN types 1-3 (CN1-CN3) are capable of eliciting the oscillatory, phasic SMP as recorded in peripheral nerves that innervate the swim appendages, the parapodia. CN1, CN2, and CN3 make monosynaptic excitatory connections onto ipsilateral, contralateral, and bilateral pedal swim-modulatory neurons [parapodial opener-phase (POP) cells], respectively; and each command cell type activates the pedal central pattern generator (CPG), leading to sustained phasic output of motor neurons and POP cells. 4. Tonic firing of CN4 causes weak activation of the SMP contralaterally. These neurons occur as two pairs of neurons in each cerebral hemiganglion, with mutual electrical and chemical synaptic interconnections. CN4 cells also excite CN1 and CN2 cells. Thus CN4 is classified as a higher-order swim command cell type. 5. Command cells classified as types 5-8 (CN5-CN8), although not capable of inducing the SMP individually, nonetheless have strong synaptic connections with pedal POP cells and/or with other command neurons. These command cells may excite or inhibit follower cells on the same or opposite sides of the preparation and modulate the swim output. 6. All the command cells tested received strong input from mechanical stimulation, either stretch or pinching, of either parapodium. Mechanosensory input from the parapodia was shown to depend on the presence of the pedal ganglion, but not the pleural. Sensory stimulation activated command cells and motor neurons, but POP cells received input from sensory stimuli only through the cerebral ganglion, probably via command cells. The effects of applied mechanosensory stimuli could be entirely mimicked by motor neuron-induced contractions of the parapodia.

Animals↗

The effects of CRF antagonists, antalarmin, CP154,526, LWH234, and R121919, in the forced swim test and on swim-induced increases in adrenocorticotropin in rats.

RATIONALE: Exposure to extreme stress has been suggested to produce long-term, detrimental alterations in the hypothalamic-pituitary-adrenal (HPA) axis leading to the development of mental disorders such as depression. Therefore, compounds that block the effects of stress hormones were investigated as potential therapeutics for depression. OBJECTIVES: In the present study, we compared the potential antidepressant-like effects of four CRF antagonists, antalarmin, CP154,526, R121919, and LWH234 (at 3, 10, and 30 mg/kg i.p., 60 min prior to the forced swim test) and the corresponding effect on swim-induced HPA activation to better elucidate the relation between HPA activity and antidepressant activity. METHODS: The antidepressant-like effects of the CRF antagonists and known antidepressants were determined in the rat forced swim test, and blood samples were obtained before and after swimming for the evaluation of adrenocorticotropin-releasing hormone (ACTH) levels. RESULTS: Antalarmin, CP154,526, and R121919 did not produce antidepressant-like effects in the forced swim test although these compounds decreased swim-induced increases in ACTH to various extents. In contrast, LWH234 reduced immobility in the forced swim test, without altering the swim-stress-induced ACTH response. However, this compound antagonized restraint-induced ACTH release. CONCLUSIONS: These data suggest that reducing stress-induced increases in HPA activity alone may not be sufficient to produce antidepressant-like activity; however, reductions in HPA activity may contribute to antidepressant actions of some treatments. In addition, it is proposed that CRF antagonists may alter differentially the HPA axis depending on the type of stressor used or behavioral measure evaluated.

Adrenocorticotropic Hormone↗

Swimming efficiency and the influence of morphology on swimming costs in fishes.

Swimming performance is considered a main character determining survival in many aquatic animals. Body morphology highly influences the energetic costs and efficiency of swimming and sets general limits on a species capacity to use habitats and foods. For two cyprinid fishes with different morphological characteristics, carp (Cyprinus carpio L.) and roach (Rutilus rutilus (L.)), optimum swimming speeds (U(mc)) as well as total and net costs of transport (COT, NCOT) were determined to evaluate differences in their swimming efficiency. Costs of transport and optimum speeds proved to be allometric functions of fish mass. NCOT was higher but U(mc) was lower in carp, indicating a lower swimming efficiency compared to roach. The differences in swimming costs are attributed to the different ecological demands of the species and could partly be explained by their morphological characteristics. Body fineness ratios were used to quantify the influence of body shape on activity costs. This factor proved to be significantly different between the species, indicating a better streamlining in roach with values closer to the optimum body form for efficient swimming. Net swimming costs were directly related to fish morphology.

Animals↗

The effect of high- and low-intensity warm-up on the physiological responses to a standardized swim and tethered swimming performance.

This investigation was conducted to determine the effect of high- and low-intensity warm-ups on physiological responses, lactate accumulation, and high-intensity freestyle and tethered swimming performance. Ten male collegiate swimmers were tested for maximal oxygen uptake (VO2 max) followed by two series of three warm-up protocols performed in a randomized order at least 2 days apart. The warm-up protocols were: (1) no warm-up (NWU), (2) a 366-m swim at 70% VO2 max (LWU) and (3) four 46-m swims at 1-min intervals at a speed corresponding to 110% VO2 max (HWU). Five minutes after each warm-up in the first series, the swimmers swam a 183-m standardized freestyle swim at a velocity corresponding to 110% VO2 max, and 5 min after each warm-up in the second series the swimmers completed a tethered swim to exhaustion with a weight attached to the tether to elicit fatigue at about 2 min. Three minutes after each warm-up and 3 min after each standardized and tethered swim, a finger-prick blood sample for lactate measurement was obtained. Heart rate and VO2 were also measured during the warm-up and the standardized and tethered swims. The performance times in the tethered swim were not significantly different between the three conditions (116.8 +/- 46.8, 137 +/- 53.3 and 122.94 +/- 37.2 s for the NWU, LWU and HWU, respectively).(ABSTRACT TRUNCATED AT 250 WORDS)

Adult↗

Synaptic basis of swim initiation in the leech. III. Synaptic effects of serotonin-containing interneurones (cells 21 and 61) on swim CPG neurones (cells 18 and 208).

Serotonin-containing cells 21 and 61 strongly excite a swim central pattern generator (CPG) neurone, cell 208, in nearby segmental ganglia in the leech Macrobdella decora. This excitatory effect is apparently independent of activity in the swim-initiating neurone cell 204, which monosynaptically excites cell 208 (Weeks, 1982b). Cell 208 excites cell 21, apparently directly. This is the first identified direct pathway for feedback from the swim central pattern generator to a swim initiator neurone. Focally applied serotonin has no effect on the soma of cell 208, but causes both excitatory and inhibitory responses in cell 208 when applied to different places within the neuropile. Cell 61 polysynaptically excites distant, posterior cells 208. This excitation is mediated at least in part by the activation of nearby cells 208, which polysynaptically excite posterior cells 208. Cell 208 is dye-coupled intraganglionically to a newly identified pair of neurones, designated cells 18. Cell 208 also excites posterior cells 18, apparently directly. This interaction may be the pathway whereby cell 61 polysynaptically excites posterior cells 208. During swimming, cell 18's membrane potential oscillates in phase with cell 208. Intracellular current injection into cell 18 during swimming perturbs the swim motor pattern. Therefore, cell 18 qualifies as a candidate swim CPG neurone.

Animals↗

The influence of temperature on muscle function in the fast swimming scup. I. Shortening velocity and muscle recruitment during swimming.

In this study, electromyography showed that scup can swim to a maximum speed of 80 cm s-1 with their red muscle whereas previous results showed that carp can swim to only 45 cm s-1. Our aim was to evaluate the adaptations that enable scup to swim nearly twice as fast as carp. Although we anticipated that, at their respective maximum speeds, the red muscle of scup would be shortening at twice the velocity (V) of carp muscle, we found that the values of V were the same (2.04 muscle lengths s-1). At any given swimming speed, V was higher in carp than in scup because carp had a larger sarcomere length excursion and higher tail-beat frequency. The smaller sarcomere excursion in scup is primarily associated with using a less undulatory style of swimming (i.e. with a smaller backbone curvature). This less undulatory style of swimming may be an important adaptation that not only reduces V but may also reduce drag. At their respective maximum speeds, however, the 28% lower sarcomere length excursion in scup is balanced by a 26% higher tail-beat frequency, giving an equal V to that of carp. Although the scup in this study were somewhat longer than the carp in the previous one (19.7 vs 13.4 cm), we believe that many of the observed differences are species-related rather than size-related. We also found that scup swam in a kinematically similar fashion at 10 degrees C and 20 degrees C. However, at 10 degrees C, the scup could swim to only 54 cm s-1 before recruiting their white muscle whereas, at 20 degrees C, they could swim to 80 cm s-1. The difference in speed of initial white muscle recruitment, as well as information on muscle mechanics, suggests that the scup compress their recruitment order into a narrow speed range at low temperatures, thereby recruiting more muscle fibres. Quantitative analysis of red muscle electromyograms in this paper supports this hypothesis.

Animals↗

Semen preparation by standard swim-up versus swim-up with test yolk buffer incubation in intrauterine insemination: a randomized study.

In order to compare the standard swim-up semen preparation with and without test yolk buffer (TYB) incubation in intrauterine insemination (IUI), we conducted a prospective multicentre randomized trial. A total of 121 infertile couples with male factor (n = 52) or unexplained infertility (n = 69) was randomly assigned to two groups following ovulation induction. Semen was prepared by standard swim-up in group A (n = 64) and by swim-up followed by TYB incubation in group B (n = 57). A maximum of two IUI cycles was performed. A total of 104 cycles was performed in the swim-up group and 90 in the TYB group. Overall, 15 pregnancies were achieved in group A and 23 in group B, with an overall pregnancy rate of 24.8 and 50.0% per patient respectively (chi2(1), P < 0.05). In the male factor group, pregnancy was achieved in six out of 24 couples (25%) following standard swim-up and in six out of 28 (21.4%) following swim-up and TYB incubation (chi2(1), not significant). In the unexplained infertility group, pregnancy was recorded in nine out of 40 couples (22.5%) following standard swim-up and in 17 out of 29 couples (58.6%) following swim-up and TYB incubation (chi2(1), P < 0.05).

Female↗

Involvement of dopamine D2 receptor mechanism in the REM sleep deprivation-induced increase in swimming activity in the forced swimming test.

Effects of monoamine synthesis inhibitors and dopamine antagonists on rapid eye movement sleep (REMs) deprivation treatment-induced increase in swimming activity were examined. Mice were deprived of REMs for 48 h by a small pedestal method. Swimming activity in REMs-deprived mice was significantly higher than those in group-housed or socially isolated animals used as the control. dl-alpha-Methyl- p-tyrosine methyl ester HCl (250 mg/kg, IP) decreased the swimming activity in REMs-deprived mice, whereas neither disulfiram (400 mg/kg, SC), a noradrenaline synthesis inhibitor, nor dl-p-chlorophenylalanine methyl ester HCl (300 mg/kg, IP) changed it. (+)-SCH23390 HCl (30 and 100 micrograms/kg, IP), a selective dopamine D1 antagonist, did not affect the activity in REMs-deprived mice. (+/-)-Sulpiride (12.5 and 25 mg/kg, IP), a selective dopamine D2 antagonist, dose-dependently decreased swimming activity in REMs-deprived mice, while it did not significantly change the swimming activity in the control animals. These results suggest that REMs deprivation treatment-induced increase in swimming activity is mainly due to the functional changes in the dopaminergic system rather than the noradrenergic or serotonergic system, and that dopamine D2 but not D1 receptor mechanism is involved in the increase in swimming activity in REMs-deprived animals.

Animals↗

Neurobehavioral studies of forced swimming: the role of learning and memory in the forced swim test.

1. Immobility in the forced swim test ("behavioral despair test") has often been regarded as an animal model of despair or depression. 2. Behavioral studies of forced swimming ("behavioral despair") are reviewed and compared with certain behavioral effects of exposure to inescapable shock (i.e., "learned helplessness"). 3. Exposure to inescapable shock clearly impairs subsequent coping responses. However, detailed behavioral studies of forced swimming indicate that immobility during forced swimming is not a failure of coping but instead reflects a relatively successful coping strategy that employs energy conserving behaviors. 4. Certain neurobiological studies of forced swimming are reinterpreted in light of the behavioral evidence that immobility during forced swimming reflects effects of learning and memory rather than effects of despair or depression. 5. Some implications for future neurobehavioral studies of forced swimming and uncontrollable shock are discussed.

Animals↗

Effects of ethanol on fight- or swim-stressed mice in Porsolt's swim test.

The effects of ethanol in Porsolt's swim test on mice preexposed to fight- or swim-stressors were investigated. The control mice did not change their behavior in the swim test after an acute injection of 0.4 or 0.8 g/kg ethanol; 1.2 g/kg ethanol increased their immobility in one but not in another experiment. The mice exposed to continuous fight-attacks in their home cage by one dominant mouse shortened immobility after 0.8 g/kg ethanol as well as tended to shorten it after 0.4 g/kg ethanol. The mice that were forced to swim in the water twice before the actual swim test responded to 0.4 g/kg ethanol by shortening immobility; 0.8 g/kg tended to have the same effect; 1.2 g/kg ethanol just failed to lengthen immobility of the fight-stressed mice and had no effect on the swim-stressed mice. Because antidepressant drugs decrease and stressors increase immobility in the swim test, the test may serve as a putative animal model of depression. The present findings showed that low doses of ethanol reverse lengthened immobility of mice preexposed to a stressor. This suggests that ethanol either has antidepressant-like properties, or it improves animal's ability to cope with a stressful situation, or both.

Aggression↗

Swimming lessons, swimming ability, and the risk of drowning.

Drowning is a leading cause of injury related death in many countries. Strategies to prevent these deaths depend upon characteristics of the victim and the specific circumstances surrounding the event. One preventive strategy that may be beneficial for persons of all ages and under nearly all circumstances is increased swimming ability, through some form of swimming instruction. However, a clear protective relationship between increased swimming ability and the risk of drowning has never been demonstrated. Studies focused on children, suggest that swimming ability may confer some protection, although the data are far from conclusive. This paper (1) reviews the current evidence regarding the relationship between swimming ability, swimming lessons and the risk of drowning, (2) reviews the past and present recommendations for swimming instruction and (3) outlines future research needs.

Adolescent↗

Bilateral inter-arm coordination in freestyle swimming: effect of skill level and swimming speed.

The aim of this study was to examine the influence of level of skill and swimming speed on inter-limb coordination of freestyle swimming movements. Five elite (2 males, 3 females; age 18.9+/-1.0 years, height 1.71+/-0.04 m, body mass 62.1+/-7.0 kg) and seven novice (age 22.0+/-2.0 years, height 1.77+/-0.04 m, body mass 74.8+/-9.0 kg) swimmers swam a sprint and a self-paced 25 m freestyle trial. The swimming trials were recorded by four digital cameras operating at 50 Hz. The digitized frames underwent a three-dimensional direct linear transformation to yield the three-dimensional endpoint kinematic trajectories. The spatio-temporal relationship between the upper limbs was quantified by means of the peak amplitude and time lag of the cross-correlation function between the right and left arm's endpoint trajectories. A strong anti-phase coupling between the two arms, as confirmed by peak amplitudes greater than 0.8, was noted for both groups and swimming speeds. Significantly higher (P<0.05) peak amplitudes were observed for the sprint compared with self-paced swimming. No significant differences in the strength of inter-limb coupling were noted between the elite and novice swimmers (P>0.05). Time lags were very close to 0 ms and did not differ between groups or swimming speeds. We conclude that in freestyle swimming, the intrinsic anti-phase (180 degrees phase difference) inter-limb relationship is strongly preserved despite the physically powerful environmental influence of the water and this "preferred" pattern is not affected by level of skill. In contrast, increasing movement speed results in stronger inter-limb coupling that is closer to the anti-phase inter-limb relationship.

Acceleration↗

Enhancement of blood lactate clearance following maximal swimming. Effect of velocity of recovery swimming.

Swimming is an endurance-intensive sport resulting in accumulation of lactate. Repeat performances are often necessary in championship events. Lactate produced during a maximal effort requires time to metabolize to a base level. If this does not occur, performance in a repeat effort may be impaired. Thus, techniques to enhance lactate clearance are of potential benefit to the athlete. We have demonstrated previously that swimming at 65% of maximum velocity significantly improved lactate clearance over passive resting. This study tested the effect of various swimming velocities on lactate clearance. Following a maximal swim, blood lactate clearance was tracked during a 15 minute cool down swim. Velocities of 55%, 65%, and 75% of maximum were tested. The results confirmed that cool down swimming will return lactate values to near resting levels in the test interval. However, statistical superiority of any of the test velocities was not demonstrated. The intensity of the swim should be below the lactate accumulation level. The 65% of maximum velocity was felt by all swimmers to be most comfortable and is a good target velocity for the athlete to reference.

Anaerobic Threshold↗

The hydrodynamics of eel swimming II. Effect of swimming speed.

Simultaneous swimming kinematics and hydrodynamics are presented for American eels, Anguilla rostrata, swimming at speeds from 0.5 to 2 L s(-1). Body outlines and particle image velocimetry (PIV) data were collected using two synchronized high-speed cameras, and an empirical relationship between swimming motions and fluid flow is described. Lateral impulse in the wake is estimated assuming that the flow field represents a slice through small core vortex rings and is shown to be significantly larger than forces estimated from the kinematics via elongated body theory (EBT) and via quasi-steady resistive drag forces. These simple kinematic models predict only 50% of the measured wake impulse, indicating that unsteady effects are important in undulatory force production. EBT does, however, correctly predict both the magnitude and time course of the power shed into the wake. Other wake flow structures are also examined relative to the swimming motions. At all speeds, the wake contains almost entirely lateral jets of fluid, separated by an unstable shear layer that rapidly breaks down into two vortices. The jet's mean velocity grows with swimming speed, but jet diameter varies only weakly with swimming speed. Instead, it follows the body wavelength, which changes more among individuals than at different speeds. Circulation of the stop-start vortex, shed each time the tail changes direction, can also be predicted at low speeds by the integral of squared tail velocity over half of a tail beat. At high speeds, these kinematics predict more circulation than is actually present in the stop-start vortex. Finally, the cost of producing the wake, one component of the total cost of transport, increases with swimming speed to the 1.48 power, lower than would be expected if the power coefficient remained constant over the speed range examined.

Anguilla↗

Effects of fin size on swimming performance, swimming behaviour and routine activity of zebrafish Danio rerio.

The zebrafish Danio rerio exhibits substantial morphological variability in the sizes and shapes of the body and the caudal fin. The present study describes swimming performance, swimming behaviour and routine locomotor activity patterns in three of the major morphotypes: wild-type, long-finned and no-tail. Wild-type and long-finned differ in total length (TL), fork length (FL), caudal fin length (CFL) and caudal fin height (CFH). No-tail has no caudal fin and is significantly smaller in standard length (SL) than the other types. Critical swimming speeds (U(crit)) were measured at 28 degrees C in a modified Brett-type water tunnel. U(crit) of wild-type fish was 56.0+/-4.8 cm s(-1) or 15.5 SL s(-)(1) (mean +/- s.d., N=21), significantly faster than the U(crit) of long-finned fish (43.7+/-6.8 cm s(-1) or 12.5 SL s(-1), N=17); both were significantly faster than the U(crit) of no-tail fish (19. 8+/-4.7 cm s(-1) or 6.9 SL s(-1), N=15). When forced to swim in the water tunnel, zebrafish tended to turn and swim downstream for short periods at slow water velocities. Turning frequencies (turns per minute, f(T)) at the slowest velocity (4 cm s(-1)) were 10. 1+/-6.5 min(-)(1) (N=63) and 8.6+/-4.7 min(-1) (N=51) for wild-type and long-finned, respectively, significantly different from that of the no-tail fish, 4.7+/-2.8 min(-1) (N=45). These frequencies decreased below 1 min(-1) at 56%, 64% and 61% of U(crit) in wild-type, long-finned and no-tail fish, respectively. Activity levels of wild-type fish were generally significantly higher than those of long-finned fish, and the levels of both were significantly higher than those of no-tail fish. The pattern of differences in relative activity levels between types was similar to that for U(crit). The results show that the wild-type fish, on a size-scaled basis, is one of the fastest-swimming fishes ever measured, reaching the maximum predicted theoretical sustained swimming speed. U(crit) of long-finned fish was 22% lower than that of wild-type fish, and U(crit) of no-tail fish was 65% lower. Similar differences were found in turning frequencies and routine activity level.

Animals↗

The respiratory metabolism of temperature-adapted flatfish at rest and during swimming activity and the use of anaerobic metabolism at moderate swimming speeds.

(1) The standard oxygen consumption and the oxygen consumption during measured swimming activity have been determined in three flatfish species at 5, 10 and 15 degrees C. (2) The relationship between weight and standard oxygen consumption for flatfish conform to the general relationship Y = aWb. On an interspecies basis, standard oxygen consumption of flatfish is significantly lower than that of roundfish. (3) A semilogarithmic model describes the relationship between oxygen consumption and swimming speed for the three species. Values for maximum oxygen consumption, metabolic scopes and critical swimming speeds are low in comparison to salmonids. (4) The optimum swimming speeds and critical swimming speeds of flatfish are similar. It is suggested that, over long distances, flatfish adopt a strategy of swimming at supercritical speeds with periods of intermittent rest to repay the accrued oxygen debt. (5) Elevated lactic acid levels in flounder white muscle after moderate swimming indicate an additional 15% anaerobic contribution to the cost of locomotion as calculated from aerobic considerations.

Acclimatization↗

Effects of swimming and land exercises versus swimming and water exercises on body composition of college students.

The purpose of this study was to examine the effects of 2 different swimming and calisthenic exercise programs on body composition and swim performance. Subjects were forty-two students (M = 19; F = 23; mean age = 20.8 yr). The experimental groups (Swim-Land [SL], n = 14; Swim-Water, [SW], n = 16) participated in 35-40 min exercise sessions which consisted of a 5-10 min warm-up, 15-20 min of swimming, and 10-15 min of calisthenics three times per week for 8 weeks. The SL experimental group performed calisthenics on land with surgical tubing while the SW experimental group performed comparable calisthenics in the water. Subjects were pretested and posttested utilizing hydrostatic weighing to determine body density and calculate percent fat. Skinfold measures and the 12-min swim for distance were also measured. Results indicated a significant 20% increase in swim performance in both experimental groups (SL = 455 +/- 144 m to 553 +/- 114 m; SW = 465 +/- 122 m to 556 +/- 123 m; p less than 0.05) but no difference between groups.(ABSTRACT TRUNCATED AT 250 WORDS)

Body Composition↗