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The activity of abdominal stretch receptors during non-giant swimming in the crayfish Cherax destructor and their role in hydrodynamic efficiency.

Recordings were made from the nerve innervating the stretch receptors of the abdominal muscle receptor organs and slow extensor muscles of tethered crayfish, Cherax destructor, during so-called "non-giant swimming". The stretch receptors were active during the flexor phase of swimming but the duration and pattern of activity varied from cycle to cycle. Their pattern of firing was modified by the activity of the large accessory neurons which make direct inhibitory synapses upon them. Neither the stretch receptors nor the accessory neurons were active during the extensor phase of the cycle. The timing and extent of tailfan movements during the period of stretch receptor activity were measured from video records before and after the stretch receptor nerves were cut in the second to fifth segments. The promotion of the tailfan during flexion was significantly delayed and the minimum angle to which the uropods were remoted at the end of flexion significantly larger in denervated animals. We propose that afferent information from the stretch receptors coordinates the timing and extent of tailfan movements according to variations in the positioning and movement of the abdominal segments such that the hydrodynamic efficiency of the tailfan is enhanced on a cycle by cycle basis during non-giant swimming.

Abdominal Muscles↗

Mechanisms of postinhibitory rebound and its modulation by serotonin in excitatory swim motor neurons of the medicinal leech.

Postinhibitory rebound (PIR) is defined as membrane depolarization occurring at the offset of a hyperpolarizing stimulus and is one of several intrinsic properties that may promote rhythmic electrical activity. PIR can be produced by several mechanisms including hyperpolarization-activated cation current (I(h)) or de-inactivation of depolarization-activated inward currents. Excitatory swim motor neurons in the leech exhibit PIR in response to injected current pulses or inhibitory synaptic input. Serotonin, a potent modulator of leech swimming behavior, increases the peak amplitude of PIR and decreases its duration, effects consistent with supporting rhythmic activity. In this study, we performed current clamp experiments on dorsal excitatory cell 3 (DE-3) and ventral excitatory cell 4 (VE-4). We found a significant difference in the shape of PIR responses expressed by these two cell types in normal saline, with DE-3 exhibiting a larger prolonged component. Exposing motor neurons to serotonin eliminated this difference. Cs+ had no effect on PIR, suggesting that I(h) plays no role. PIR was suppressed completely when low Na+ solution was combined with Ca2+-channel blockers. Our data support the hypothesis that PIR in swim motor neurons is produced by a combination of low-threshold Na+ and Ca2+ currents that begin to activate near -60 mV.

Animals↗

Energy metabolism of underwater swimming in river-otters (Lutra lutra L.).

We used a still-water swim channel in conjunction with open-flow oxygen and carbon dioxide respirometry to examine the energy requirements of river-otters (Lutra lutra L.) swimming voluntarily underwater in Neumünster Zoo (Germany). While at rest on land (5 degrees C), river-otters had a respiratory quotient of 0.77 and a resting metabolic rate of 4.1 W kg-1. This increased to an estimated 6.4 W kg-1 during rest in water (11-15 degrees C) and to 12.3 W kg-1 when the animals were feeding in the channel. River-otters swimming under water preferred a mean speed of 0.89 m s-1, and their energy requirements attained 11.6 W kg-1. Cost of transport, however, was minimal at 1.3 m s-1 and amounted to 0.95 J N-1 m-1.

Animal Welfare↗

An integrative study of the temperature dependence of whole animal and muscle performance during jumping and swimming in the frog Rana temporaria.

The aims of this study were: (1) to analyze individual variation in frog locomotor performance, (2) to compare the thermal sensitivity of jumping and swimming, and (3) to contrast whole animal versus muscle fiber performance at different temperatures. The jumping and swimming performance of Rana temporaria was analyzed at 5, 10, 15 and 20 degrees C. Muscle fiber bundles were isolated from lateral gastrocnemius and subjected to the length and activation patterns thought to occur in vivo. As temperature increased, locomotor performance in R. temporaria improved with a Q10 of 1.2 for both jump take-off velocity and mean swimming velocity. The slope of the relationship between performance and temperature (TE) was similar for both locomotor parameters and was described by the equation z-scores of locomotor performance = 0.127 x TE - 1.585. Although some frogs performed better than others relative performance was affected by locomotor type and temperature. Locomotor performance improved with temperature as the power required during take-off and the mean muscle power output increased with Q10 values of 1.7 and 1.6 respectively. The mean muscle power output during take-off was only 34% of the calculated requirements for the whole animal, suggesting the involvement of elastic strain energy storage mechanisms.

Animals↗

Passive and active floating torque during swimming.

The purpose of this study was to examine the effect of passive underwater torque on active body angle with the horizontal during front crawl swimming and to assess the effect of body size on passive torque and active body angle. Additionally, the effects of passive torque, body angle and hydrostatic lift on maximal sprinting performance were addressed. Ten boys [aged 11.7 (0.8) years] and 12 male adult [aged 21.4 (3.7) years] swimmers volunteered to participate. Their body angle with the horizontal was measured at maximal velocity, and at two submaximal velocities using an underwater video camera system. Passive torque and hydrostatic lift were measured during an underwater weighing procedure, and the center of mass and center of volume were determined. The results showed that passive torque correlated significantly with the body angle at a velocity 63% of v(max) ( alpha(63) r=-0.57), and that size-normalized passive torque correlated significantly with the alpha(63) and alpha(77) (77% of v(max)) with r=-0.59 and r=-0.54 respectively. Hydrostatic lift correlated with alpha(63) with r=-0.45. The negative correlation coefficients are suggested to be due to the adults having learned to overcome passive torque when swimming at submaximal velocities by correcting their body angle. It is concluded that at higher velocities the passive torque and hydrostatic lift do not influence body angle during swimming. At a velocity of 63% of v(max), hydrostatic lift and passive torque influences body angle. Passive torque and size-normalized passive torque increases with body size. When corrected for body size, hydrostatic lift and passive torque did not influence the maximal sprinting velocity.

Adult↗

Relative contribution of arms and legs in humans to propulsion in 25-m sprint front-crawl swimming.

Eight male subjects were asked to swim 25 m at maximal velocity while the use of the arm(s) and legs was alternately restricted. Four situations were examined using one arm (1A), two arms (2A), one arm and two legs (1A2L) and both arms and legs (2A2L, normal swim) for propulsion. A significant mean increase of 10% on maximal velocity was obtained in 1A2L and 2A2L compared to 1A and 2A. A non-significant 4% effect was obtained in 1A. This study focused on the actual contribution of leg kick in the 10% gain in maximal velocity. It was clear that the underwater trajectory of the wrist was modified by the action of the legs (most comparisons P < 0.001). Therefore it was thought that the legs enhanced the generated propulsive force by improving the propulsive action of the arm. The arm action was quantified by selecting typical phases from the filmed trajectory of the wrist, namely forward (F), downwards (D) and backwards (B). Although there was a tendency for individual changes in kinematic parameters (F, D and B) to occur with individual changes in velocity when 2A was compared to 2A2L, no relationship was found between the relative changes in F, D and B and relative changes in velocity. This was illustrated by describing the responses of three individuals who could represent three patterns of contribution by legs and arms to propulsion in high speed swimming.

Arm↗

Dose-dependent effects of noradrenergic denervation by DSP-4 treatment on forced swimming and beta-adrenoceptor binding in the rat.

DSP-4 is a neurotoxin highly selective for the noradrenergic nerve terminals originating from the locus coeruleus. Preliminary data suggested that its effect in a typical screening test for antidepressant drugs, the forced swimming test, is biphasic dependent on the dose. In the present study, DSP-4 was administered in four doses (5, 10, 30 and 50mg/kg) to male Wistar rats. Administration of the neurotoxin had a dose-dependent biphasic effect on immobility time in the forced swimming test 8 and 9 days later. Thus, DSP-4 at the dose of 10mg/kg increased immobility, but higher doses reduced this measure. The reduction of noradrenaline concentration in the frontal cortex and hippocampus was dose-dependent starting from the dose 10mg/kg. Cortical beta-adrenoceptor binding was increased by DSP-4 treatment at the doses 30mg/kg and 50mg/kg. These results suggest that the increase in immobility time in the forced swimming test is associated with presynaptic changes in noradrenaline availability, whereas the decrease in immobility observed after more complete denervation is associated with postsynaptic receptor supersensitivity.

Adrenergic Agents↗

Recruitment patterns in the rat hindlimb muscle during swimming.

Cycle periods and electromyographic (EMG) burst durations of selected extensors and flexors are shorter during swimming than during running at 27 m/min, while the relative ON-OFF timing of these interrelationships are similar. The mean EMG activities of the tibialis anterior and adductor longus were higher and the soleus and medial gastrocnemius were lower during swimming than during running. Both fast and slow extensors are activated during both forms of locomotion, thus demonstrating that there is not a selective recruitment of fast or selective inhibition of slow ankle extensors during swimming as measures of muscle blood flow have suggested.

Animals↗

Opioid and non-opioid stress analgesia from cold water swim: importance of stress severity.

We have previously reported that stress severity plays an important role in determining the neurochemical basis of stress-induced analgesia from inescapable footshock. Increasing severity (duration or intensity of continuous footshock) causes a shift in mediation of the resultant analgesia from opioid to non-opioid. In this study, we find that stress severity plays a similar role in analgesia from cold water swim. More severe swims (longer duration or lower water temperature) produce stress analgesia insensitive to the opiate antagonist, naltrexone, whereas less severe swims produce analgesia significantly attenuated by this drug.

Analgesia↗

Strain differences in the magnitude of swimming-induced analgesia in mice correlate with brain opiate receptor concentration.

Swimming-induced analgesia was studied in 4 strains of mice differing in central opiate receptor density: C57BL/6By (C57), BALB/cBy (BALB/c), CXBK and CXBH. The degree of 'swim analgesia' significantly differed among strains in the order CXBH greater than BALB/c = C57 greater than CXBK. This order positively correlates with known differences in opiate receptor density in these strains. Naloxone reversed the analgesic effect of swimming in CXBH, C57 and BALB/c, but was ineffective in opiate receptor-deficient CXBK mice. These results suggest that genetic differences in central opiate receptor density influence the analgesic response to stressful stimuli.

Analgesia↗

Motoneuronal commands during swimming behaviour in the shore crab.

Neurograms of proximal leg motor nerves were obtained during swimming in the shore crab. Whereas excitor motoneurones fire in bursts, the common inhibitor motoneurone discharges tonically with simultaneous spikes in all the motor nerves. The average firing frequency of the common inhibitor increases as the period of the swimming cycle decreases. Moreover, greater fluctuations of the firing frequency of the common inhibitor occurs within long rather than short swim cycle periods.

Action Potentials↗

Hippocampal RSA (theta), apnea, bradycardia and effects of atropine during underwater swimming in the rat.

Hippocampal RSA (theta), neocortical EEG, and heart rate were recorded during surface and underwater swimming in the rat. RSA was present with slightly reduced frequency during the apnea, bradycardia, and vibrissae arrest associated with under water swimming. Atropine sulfate (50 mg/kg) blocked bradycardia did not affect RSA. Contrary to previous suggestions, no causal relation was found between RSA and respiration, heart rate, or vibrissae movement. The study supports the view that RSA is related to the neural control of voluntary movement. It is suggested that the slight reduction in RSA frequency during underwater swimming may be part of an oxygen conservation process.

Animals↗

The effect of gonadectomy on antinociception induced in mice by swim stress.

A 30 s swim in water at 30 degrees C reduced the number of abdominal constrictions produced in mice by i.p. acetic acid. Naloxone antagonised this stress-induced antinociception in the female but not in the male. Orchidectomy altered neither the antinociception nor its insensitivity to naloxone antagonism. However, oophorectomy completely abolished the antinociception induced by the swim. These results suggest a sex difference in the involvement of the endogenous opioid system in swim stress-induced antinociception in mice.

Animals↗

8-Hydroxy-2-(di-n-propylamino)tetralin, a selective serotonin1A receptor agonist, reduces the immobility of rats in the forced swimming test by acting on the nucleus raphe dorsalis.

8-Hydroxy-2-(di-n-propylamino)tetralin (8-OH-DPAT), a selective serotonin1A (5-HT1A) receptor agonist, was studied for its anti-immobility activity in the forced swimming test when administered into the raphe nuclei medianus and dorsalis of rats. At concentrations ranging from 0.5 to 5 micrograms, 8-OH-DPAT significantly reduced the immobility of rats when administered into the nucleus raphe dorsalis, but only 5 micrograms was effective when administered into the nucleus medianus. The activity of rats in an open-field under conditions identical to those used in the forced swimming test was not significantly changed by various concentrations of 8-OH-DPAT administered into the nucleus raphe dorsalis, but was significantly increased by an infusion of 5 micrograms 8-OH-DPAT into the nucleus raphe medianus. The effect of an infusion of 1 micrograms 8-OH-DPAT into the nucleus dorsalis was prevented by infusing 2.5 micrograms (-)-propranolol or 2.5 micrograms (-)-pindolol into the same area 5 min before 8-OH-DPAT or by treating the animals with sulpiride systemically (100 mg/kg i.p.) or centrally (in the nucleus accumbens; 1 microgram/0.5 microliter). The results suggest that 8-OH-DPAT reduces the immobility of rats by activating dopamine transmission, probably in the nucleus accumbens, as a consequence of its ability to reduce the activity of 5-HT neurons that originate in the nucleus raphe dorsalis. In view of the similarities between the effects of well-established antidepressants and 8-OH-DPAT in the forced swimming test, it is suggested that 5-HT1A receptor agonists may constitute a novel class of antidepressant agents.

8-Hydroxy-2-(di-n-propylamino)tetralin↗

Morphine analgesia and tolerance in mice selectively bred for divergent swim stress-induced analgesia.

Morphine-induced analgesia and tolerance were examined in Swiss Webster mice selectively bred for high and low swim stress-induced analgesia. Morphine produced a dose-dependent analgesia in both lines; it was 4-fold more potent in the high analgesia line than in the low analgesia line. Despite the differences in morphine-induced analgesia, the degree of tolerance was the same in both lines. Together, these data suggest that selective breeding of mice for high and low swim stress-induced analgesia produced a striking difference in morphine-induced analgesia without affecting the degree of tolerance. Thus, while there is a common genetic determination in swim stress-induced and morphine-induced analgesia, the development of tolerance to morphine possibly relies on a different genetic background.

Analgesia↗

Effects of social isolation on brain catecholamines and forced swimming in rats: prevention by antidepressant treatment.

Post-weaning rats were housed alone or in groups for a period of 4 or 8 weeks. A portion of the animals received tricyclic antidepressant treatment, desipramine 20 mg/kg/day, during this period. Animals were then tested behaviorally by forced swimming. Isolation was associated with significantly longer durations of immobility during forced swimming. This was blocked by desipramine treatment. Desipramine treatment did not have a significant effect on the swimming durations of group-housed rats. Hindbrain and midbrain levels of catecholamines were subsequently measured and turnover rates estimated by administration of alpha-methyl-p-tyrosine or saline. Isolated rats had increased levels and decreased turnover of catecholamines. The increase in norepinephrine but not dopamine levels was blocked by desipramine, while antidepressant effects on turnover could not be tested with this method. Reduced social stimulation thus appears to be associated with reduced catecholamine release which may result in the accumulation of these transmitters in the central nervous system. Treatment with desipramine appeared essentially to compensate for reduced social stimulation, blocking isolation-induced noradrenergic neurochemical changes, while having few significant effects on control animals. This study may be helpful in furthering our understanding of how the interaction of organisms with their environment influence catecholamine systems and how antidepressants may act to restore function.

Animals↗

Reduction in cervical probing analgesia by repeated prior exposure to cold-water swims.

Previous work has established that stimulation of the vaginal cervix (cervical probing: CP) and cold-water swims (CWS) each produce analgesia in rats that is affected similarly by catecholaminergic mechanisms, but differentially by serotoninergic and opioid mechanisms. The present study examined whether CP analgesia, like CWS analgesia, would be reduced by prior repeated CWS exposure. Tail-flick latencies were significantly increased by CP and CWS in naive animals. In animals chronically preexposed to 14 daily swims, CP analgesia was reduced and CWS analgesia was reduced at 60, but not 30 min after the swim. After 1 month of recovery, CP analgesia showed a persistent reduction, whereas CWS analgesia was fully recovered. These data suggest that the mechanisms underlying CP and CWS analgesia are interrelated.

Afferent Pathways↗

Age-related decrements in the analgesic response to cold-water swims.

Among the compromised physiological responses affected by aging is an impaired ability to thermoregulate in a cold environment. Since acute exposure to cold-water swims (CWS) produces both analgesic and hypothermic responses in young adult rats, the present study examined whether systematic variations in these responses occurred as a function of age. Separate cohorts of 4, 9, 14, 19 and 24-month old female rats received a no-swim condition and a 2 degrees C swim for 3.5 min with tail-flick latencies, jump thresholds and core body temperature assessed 30, 60 and 90 min later. The order of conditions was counterbalanced with an interval of four days between conditions. While the four younger cohorts displayed similar CWS analgesia on the tail-flick test, the 24-month cohort failed to display CWS analgesia on this measure. Age-related differences in CWS analgesia on the jump test appeared more gradual with the three older cohorts displaying significant attenuations relative to the 9-month group, and the oldest cohort displaying significant attenuations relative to the 4-month group. In contrast, the hypothermic effect of CWS was significantly potentiated in the three oldest cohorts, with a progressive inability to cope with thermoregulation observed as a function of age. The observed decrements in the analgesic response to CWS as a function of age appear to represent a change in (a) the ability of the animal to perceive the CWS stimulus as stressful (b) an endogenous pain-inhibitory system and/or (c) an endogenous pain transmission system that is independent of thermoregulatory mechanisms.

Adrenalectomy↗