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Transformations in flagellar structure of Rhodobacter sphaeroides and possible relationship to changes in swimming speed.

Rhodobacter sphaeroides is a photosynthetic bacterium which swims by rotating a single flagellum in one direction, periodically stopping, and reorienting during these stops. Free-swimming R. sphaeroides was examined by both differential interference contrast (DIC) microscopy, which allows the flagella of swimming cells to be seen in vivo, and tracking microscopy, which tracks swimming patterns in three dimensions. DIC microscopy showed that when rotation stopped, the helical flagellum relaxed into a high-amplitude, short-wavelength coiled form, confirming previous observations. However, DIC microscopy also revealed that the coiled filament could rotate slowly, reorienting the cell before a transition back to the functional helix. The time taken to reform a functional helix depended on the rate of rotation of the helix and the length of the filament. In addition to these coiled and helical forms, a third conformation was observed: a rapidly rotating, apparently straight form. This form took shape from the cell body out and was seen to form directly from flagella that were initially in either the coiled or the helical conformation. This form was always significantly longer than the coiled or helical form from which it was derived. The resolution of DIC microscopy made it impossible to identify whether this form was genuinely in a straight conformation or was a low-amplitude, long-wavelength helix. Examination of the three-dimensional swimming pattern showed that R. sphaeroides changed speed while swimming, sometimes doubling the swimming speed between stops. The rate of acceleration out of stops was also variable. The transformations in waveform are assumed to be torsionally driven and may be related to the changes in speed measured in free-swimming cells. The roles of and mechanisms that may be involved in the transformations of filament conformations and changes in swimming speed are discussed.

Acceleration↗

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

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

Journal Article↗

Kinematics and critical swimming speed of juvenile scalloped hammerhead sharks

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

Journal Article↗

Hopping and swimming in the leopard frog, Rana pipiens: II. A comparison of muscle activities.

Electromyography (EMG) was used to examine muscle activity of the major hip, knee, and ankle extensors during both hopping and swimming in leopard frogs. Chronic EMG electrodes were implanted for periods of 7-10 days. This permitted us to record EMG activities during both hopping and swimming from the same electrode, allowing a direct comparison of the timing and amplitudes of muscle activity between the two behaviors. We could then relate these activities to the kinematics of locomotion. In both behaviors, all three extensors were synchronously activated 30-50 ms before limb extension began. However, the hip extensor turned on relatively earlier in hopping than in swimming when on time was expressed as percent of stride. The hip and knee extensors were activated relatively longer in hopping and the ankle extensor relatively longer in swimming. The amplitudes of the rectified, integrated EMG signals were roughly twice as large in hopping as in swimming for all three muscles, supporting the notion that propulsion in hopping requires more force than in swimming. The EMG burst durations differed little between the muscles or, in relative duration, between the behaviors. As has been found in other quadrupeds, the EMG bursts began before visible movement and ceased at or before hindlimb extension was completed. In our animals, however, we found a consistent, low level (10-30% of maximum amplitude) of EMG activity that continued 60-200 ms past the end of the burst and into the suspension periods in both hopping and swimming. We hypothesize that this unusual activity may be present in frogs so that the hind limb remains aero(hydro)dynamically stable as the frog arches through its leap or glides in swimming following completed limb extension. Thus, the timing and pattern of the EMG bursts are consistent with those present in other tetrapods and support conservatism of neural control. However, the prolonged low-level activity suggests flexibility in the control pattern and variation according to specific behaviors.

Animals↗

Spastic mutant axolotl: identification of a phenocopy pathway with implications for the control of axolotl swimming by the vestibulocerebellum.

The spastic mutant axolotl shows abnormal swimming behavior, which includes a preponderance of "embryonic" swimming elements (coils) versus mature swimming elements (sinusoids) and a failure to entrain sinusoids into a prolonged swimming sequence. The mutant also shows anatomical disorganization in the area acousticolateralis and cerebellar auricle, but it is unclear (1) to what extent the behavioral abnormalities are traceable to the vestibulocerebellar defect or (2) how the vestibulocerebellar pathway modulates swimming behavior in the normal axolotl. We have performed quantitative cine analysis of electric shock-induced swimming bouts in normal axolotls, spastic mutants, and a variety of neurosurgically altered wild-type axolotls. We scored the incidence of coil elements (25% in controls, 70-90% in spastics) versus sinusoid elements, as well as length distributions of coilfree intervals (short to long trains of sinusoidal swimming) and of sinusoidfree intervals (of brief of prolonged coiling). We found that bilateral VIIIth nerve lesions or surgical undercutting of the cerebellar auricle in wild-type axolotls almost exactly reproduced the behavioral deficit seen in spastic (75-81% coils, loss of long sinusoid trains, and appearance of prolonged coiling intervals at least some of which coupled several coils into trains of thrashing behavior). By contrast, neither complete transection of the CNS at low midbrain levels nor section of cranial nerves V, VII, or X (lateral line) resulted in an increased incidence of coil elements beyond 26% nor significantly altered the length distributions of S-intervals and C-intervals. Nor did any of the latter lesions disrupt the spasticlike swimming patterns of axolotls already subjected to auricle or VIIIth nerve lesions.(ABSTRACT TRUNCATED AT 250 WORDS)

Ambystoma↗

Control of leech swimming activity by the cephalic ganglia.

We investigated the role played by the cephalic nervous system in the control of swimming activity in the leech, Hirudo medicinalis, by comparing swimming activity in isolated leech nerve cords that included the head ganglia (supra- and subesophageal ganglia) with swimming activity in nerve cords from which these ganglia were removed. We found that the presence of these cephalic ganglia had an inhibitory influence on the reliability with which stimulation of peripheral (DP) nerves and intracellular stimulation of swim-initiating neurons initiated and maintained swimming activity. In addition, swimming activity recorded from both oscillator and motor neurons in preparations that included head ganglia frequently exhibited irregular bursting patterns consisting of missed, weak, or sustained bursts. Removal of the two head ganglia as well as the first segmental ganglion eliminated this irregular activity pattern. We also identified a pair of rhythmically active interneurons, SRN1, in the subesophageal ganglion that, when depolarized, could reset the swimming rhythm. Thus the cephalic ganglia and first segmental ganglion of the leech nerve cord are capable of exerting a tonic inhibitory influence as well as a modulatory effect on swimming activity in the segmental nerve cord.

Action Potentials↗

Comparison of tetrahydroaminoacridine and physostigmine on scopolamine-induced free swim behavior in the rat.

The effect of acetylcholinesterase inhibitors on free swim behavior in rats pretreated with scopolamine (0.32 mg/kg, IP) was examined. Long-Evans rats received a single 5-min testing trial in a 1.5 m black swimming pool, and swim distance in three concentric annulus corridors (peripheral, middle, and inner) and the number of body-turn transitions (greater than 45 degrees) were measured. Physostigmine (1.0 mg/kg, IP) increased swim distance in the middle and inner annulus corridors, compared to tetrahydroaminoacridine (2.0 mg/kg and 10 mg/kg, IP) and scopolamine alone (control) (Ps less than 0.01), and increased body-turn transitions, compared to all the other groups (Ps less than 0.05), but had no significant effect on peripheral annulus corridor swim distance, total swim distance, or swim speed. The results suggest that physostigmine produces uniquely different free swim patterns from tetrahydroaminoacridine following cholinergic blockade. These findings have implications for investigations attempting to restore spatial learning and navigation (e.g., Morris water maze) using acetylcholinesterase inhibitors following experimentally-induced cholinergic losses.

Animals↗

The activity of pramipexole in the mouse forced swim test is mediated by D2 rather than D3 receptors.

RATIONALE: Recent studies have reported antidepressant-like activities of the dopamine D2/D3 agonist pramipexole in the chronic mild stress model and in the forced swim test, suggesting that D3 receptor agonists may represent a new class of antidepressant drugs. However, the relative contribution of D2 or D3 receptors to the activity of pramipexole in these models is unclear. OBJECTIVES: The aim of the current studies was to explore the role of dopamine D2 and D3 receptors in the activity of pramipexole in the mouse forced swim test. METHODS: The effect of pramipexole (0.1-3.2 mg/kg) in the mouse forced swim test was examined both in conjunction with D2 and D3 receptor antagonists (haloperidol (0.1-1 mg/kg) and LU-201640 (A-437203, 5.6-17.8 mg/kg), as well as in D3 receptor knockout mice obtained on two different background strains (C57BL/6J and B6129SF2/J). Locomotor activity was also assessed following pramipexole administration. RESULTS. Pramipexole produced dose-dependent reductions in immobility in the forced swim test at doses that did not produce generalized increases in locomotor activity. LU-201640, the D3 selective antagonist, failed to block the antidepressant-like effects of pramipexole. In contrast, the efficacy of pramipexole in the forced swim test was completely blocked by the D2 antagonist, haloperidol. No baseline differences were observed between knockout and wild-type mice from either background strain in locomotor activity or in the forced swim test. Furthermore, in both background strains, pramipexole showed similar efficacy in the forced swim test for both wild-type and knockout mice. CONCLUSIONS: Taken together, these studies suggest that the D2 receptor rather than the D3 receptor is important for the antidepressant-like activity observed for pramipexole in the mouse forced swim test.

Animals↗

A selective test for antidepressant treatments using rats bred for stress-induced reduction of motor activity in the swim test.

RATIONALE AND OBJECTIVE: This paper describes a new procedure for detecting effective antidepressant treatments. The procedure uses the swim-test susceptible (Susceptible) rat which has been selectively bred to show decreased struggling behavior in a swim test after exposure to a mild stressor. The ability of treatments to block this decrease in swim-test activity was assessed as a method for detecting effective antidepressants. RESULTS: In both male and female Susceptible rats, chronic (14-day) treatment with different antidepressant drugs delivered via osmotic minipump [i.e., three tricyclics (desmethylimipramine, imipramine, amitriptyline), two selective serotonin reuptake inhibitors (fluoxetine and sertraline), a monoamine oxidase inhibitor (phenelzine), and two atypical antidepressants (venlafaxine and bupropion)] all prevented the stress-induced decrease in swim-test struggling normally shown by these rats. Electroconvulsive shock had a similar effect. Unlike antidepressant drugs, 14-day treatment with various non-antidepressant drugs [i.e., a stimulant (amphetamine), an anxiolytic (chlordiazepoxide), an antihistamine (chlorpheniramine), and an anticholinergic (scopolamine)] did not have this effect. Antidepressant drug treatment for 1 day (i.e., acute treatment) was also ineffective in this test. The procedure described above requires use of the Susceptible rat--swim test resistant rats (i.e., rats selectively bred to be resistant to decreased swim-test activity after exposure to stressful conditions) showed no significant differences in swim-test behavior between stress and nonstress conditions after 14-day drug treatment, and randomly bred Sprague-Dawley rats did not show a decrease in swim-test activity following exposure to the mild stressor that is the basis for the test. CONCLUSION: These results suggest that the procedure described here, which uses a rat subject that has been bred for vulnerability to stressful conditions, may be a selective screening technique for effective antidepressant treatments.

Animals↗

Adaptive variations of undulatory behaviors in larval lamprey: comparison of swimming and burrowing.

In larval lamprey, movements and muscle activity during swimming and burrowing behaviors were compared. Burrowing consisted of two components: an initial component in which the head was driven into the burrowing medium; and a final component in which the animal pulled the rest of its body into the burrowing medium. The initial component of burrowing was characterized by large undulatory movements and rhythmic muscle burst activity that were similar in form to those during fast swimming, but more intense. During the initial component of burrrowing, burst durations, burst amplitudes, and burst proportions of motor activity were larger than those during swimming, while cycle time was slightly shorter than during swimming. Intersegmental phase lags and right-left phase values were similar for swimming and initial burrowing. The final component of burrowing was characterized by sharp, long-duration flexures on one side of the body, sometimes followed by similar flexures on the other side. Each flexure was produced by long-duration, large-amplitude muscle burst activity on the same side of the body or several shorter sequential bursts with slightly smaller amplitudes. During the final component of burrowing, burst durations and burst amplitudes of motor activity were much larger than those during swimming or during the initial component of burrowing. It is suggested that the motor patterns for swimming and the initial component of burrowing are produced by a common spinal locomotor network. The final component of burrowing may use some of the same neurons in the spinal locomotor networks, but the networks are probably configured differently than the situation during swimming.

Adaptation, Physiological↗

Factors affecting swimming economy in children and adults.

The aim of this study was to examine the influence of several explanatory factors: anthropometry, buoyancy, passive underwater torque, drag and swimming technique on the energy cost of swimming front crawl in children and adults. Submaximal V(.)O(2) was measured in ten children (age 12) and 13 adults (age 21), as well as body length (BL), body mass, arm length, propelling size, active drag, hydrostatic lift, passive torque, intracyclic velocity fluctuation, hand slip, stroke length and body angle. The results show that body length ( r=0.74), body mass ( r=0.86) propelling size ( r=0.61), arm length ( r=0.66), distance between the center of mass and the center of volume (Delta d, r=0.74) and body angle during swimming ( r=-0.56) all showed significant linear relationships with the cost of swimming at 1.0 m x s(-1) (CS(1.0)). When normalizing the cost of swimming to body size (CS(1.0) x BL(-1)) there were no differences between the two groups. The conclusions of this study are that the combination of BL, body mass, active drag factor, passive torque, drag efficiency and hydrostatic lift were able to explain 97% of the variation in the cost of swimming for the whole group of swimmers. The size-independent factors of torque and floating abilities (density and Delta d in % of BL), together with swimming technique and active drag were found to explain 75% of the variations in CS(1.0) x BL(-1). The identical values for CS(1.0) x BL(-1) for children and adults are explained through the opposing effects of a better swimming technique in the adults, and a better passive torque in the children.

Adolescent↗

Antioxidant status in various tissues of the mouse after fasting and swimming stress.

We studied the effect of fasting and swimming stress on a number of non-enzymatic and enzymatic antioxidant factors in various mouse tissues in order to see if their action was synergic. We examined levels of reduced (GSH), oxidized (GSSG) and total glutathione, total SH groups (TSH), sum of GSH and protein sulphydryl groups of cytosolic fractions, and the activities of superoxide dismutase (SOD), catalase, glutathione peroxidase, glutathione reductase in adductor muscle, heart and liver. We also studied blood levels of GSH and glutathione bound to protein by mixed disulphides (GSSP). The case series consisted of four groups of animals (n = 10 for each group), namely no swimming and no fast, no swimming and fast, swimming and no fast, and swimming and fast. Fasting (18 h) resulted in a significant GSH depletion in all of the organs studied (-39% in the liver, -30% in the adductor muscle, -21% in the heart); GSSG increased significantly in the heart (+19%). Swimming to exhaustion, which lasted 3.95 (0.18) min [mean (SD), n = 10] with no significant difference between fast and no fast, resulted in a significant GSH depletion, to a percentage lower than that observed after fasting, in the adductor muscle and heart (-12% and -11%, respectively). In the blood of swimming mice, significant increases in GSH (+10%) and GSSG (+21%) levels were observed, whereas GSSP decreased (-15%). Enzyme activities after swimming were modified in only a few cases, and in a complex way. The findings of GSH depletion and a decrease in SOD activity in the adductor muscle seems to confirm the sensitivity of this organ to an overproduction of reactive oxygen species. At the same time, the GSSP decrease observed in blood was a new and unexpected finding, one that indicates a very prompt adaptation of red cells to increased oxidant states.

Animals↗

2-Amino-7-phosphonoheptanoic acid, a selective N-methyl-D-aspartate antagonist, blocks swim-induced elevation of cerebellar cyclic guanosine monophosphate.

In order to explore how rapidly locomotor activity induces an elevation in cerebellar cyclic guanosine monophosphate (cGMP) content, Sprague-Dawley rats, pretrained to swim a 2.5-m course, were required to swim from one to 5 laps representing from 7 to 40s of strenuous activity. Immediately after completing the swimming task, each animal was killed by microwave irradiation and the cerebellum was collected for subsequent determination of the cGMP content. There was no difference in the cerebellar cGMP content between rats swimming one lap, i.e. for 7 s, and control rats that did not swim. However, there was a linear increase in the cGMP over control values from 1.8- to 2.4-fold in rats swimming 3 and 5 times, respectively. The first significant elevation of the cerebellar cGMP was seen at 24 s (3 laps). To determine if acidic amino acid pathways were involved in this elevation, a low dosage of a selective NMDA antagonist, 2-amino-7-phosphonoheptanoic acid (APH) was injected intracerebroventricularly 4 min before having rats swim 4 laps. This low dosage of APH, which alone had no effect on the cerebellar cGMP content, completely blocked the swim-induced elevation of this parameter. These data provide the first report of how quickly locomotor activity elevates the cerebellar cGMP content and further suggest that an NMDA receptor-mediated pathway is involved in the activity-induced elevation of this parameter.

2-Amino-5-phosphonovalerate↗

Control of physical exercise of rats in a swimming basin.

To study the mutual interaction between physical exercise and antioxidant systems in rats, we selected swimming as a model for exercise performance. Swimming belongs to the natural behavior of a rat, which under proper experimental conditions, primarily involves physical exercise with little emotional arousal. Therefore, we developed a swimming basin in which the intensity of exercise was manipulated by swimming speed and swimming duration. A laser beam interruption system enables recording of swimming patterns. For comparison we also used the basin to induce emotional arousal. Hereto the basin was transformed into a maze, in which unexpected blockade of a learned swimming route induced a panic-like emotional reaction. The antioxidant enzyme superoxide dismutase decreased in rat plasma after emotional arousal, not after physical exercise. Depletion of the antioxidant glutathione in the liver by diethyl maleate led to decrease of swimming performance. Noradrenaline but not adrenaline plasma levels increased in response to physical exercise. After emotional arousal the ratio noradrenaline/adrenaline did not change. In contrast, lactate only increased in response to emotional arousal. Plasma levels of glucose increased after both stress situations. Beta-adrenoceptor function, determined in the heart and in erythrocytes, only changed after physical exercise. The sensitivity to the beta-agonist (-)isoprenaline in the right atrium decreased and a downregulation of the beta-adrenoceptor density was observed in the erythrocyte.

Animals↗

Lateralised swimming behaviour in the California sea lion.

Lateralised motor behaviour in the pinnipeds has been subject to little investigation. This study examined the swimming behaviour of seven zoo-housed California sea lions to determine whether they exhibited a directional bias in their motor behaviour. Data were collected on the direction of the animals' swimming patterns from the point of entering a pool of water from dry land. Each animal was studied for 100 episodes of swimming. All seven of the sea lions showed significant (P<0.001) bias in the direction of their swimming, although unidirectional bias was not observed at the level of the population. The direction of the sea lions' swimming patterns varied significantly according to the animals' sex. Males showed a preference at the level of the population for swimming in a clockwise direction, while females showed a population-level counterclockwise swimming preference. Overall, the findings appear to suggest that California sea lions, like other marine mammals, exhibit motor bias in the direction of their swimming patterns, although further work using larger sample sizes is needed before more firm conclusions regarding motor laterality in this species can be reached.

Animals↗

Comparison of clinical and genetic variables of cardiac events associated with loud noise versus swimming among subjects with the long QT syndrome.

Acute auditory stimuli and swimming activities are frequently associated with syncope, aborted cardiac arrest, and death in the long QT syndrome (LQTS). We investigated the clinical and genetic findings associated with cardiac events precipitated by these arousal factors. The study population involved 195 patients with an index cardiac event associated with a loud noise (n = 77) or swimming activity (n = 118). Patients with events associated with loud auditory stimuli were older at their index event and were more likely to be women than patients who experienced events during swimming-related activities. Patients with an index event associated with loud noise were likely to have subsequent events related to auditory stimuli; patients with an index event associated with swimming were likely to have recurrent events related to swimming or physical activities. Family patterning of auditory and swimming and/or physical activity-related events was evident. Genotype analyses in 25 patients revealed a significant difference in the distribution of index cardiac events by genotype (p <0.001), with all 19 patients with swimming-related episodes associated with LQT1 genotype and 5 of 6 patients with auditory-related events associated with LQT2 genotype. The clinical profile and genotype findings of patients with LQTS who experience cardiac events related to acute auditory stimuli are quite different from those who experience events accompanying swimming activities.

Acoustic Stimulation↗

Occupational exposure to trihalomethanes in indoor swimming pools.

The study evaluated occupational exposure to trihalomethanes (THMs) in indoor swimming pools. Thirty-two subjects, representing the whole workforce employed in the five public indoor swimming pools in the city of Modena (Northern Italy) were enrolled. Both environmental and biological monitoring of THMs exposure were performed. Environmental concentrations of THMs in different areas inside the swimming pools (at the poolside, in the reception area and in the engine-room) were measured as external exposure index, while individual exposure of swimming pool employees was estimated by THMs concentration in alveolar air. The levels of THMs observed in swimming pool water ranged from 17.8 to 70.8 microg/l; the mean levels of THMs in ambient air were 25.6+/-24.5 microg/m3 in the engine room, 26.1+/-24.3 microg/m3 in the reception area and 58.0+/-22.1 microg/m3 at the poolside. Among THMs, only chloroform and bromodichloromethane were always measured in ambient air, while dibromochloromethane was detected in ambient air rarely and bromoform only once. Biological monitoring results showed a THMs mean value of 20.9+/-15.6 microg/m3. Statistically significant differences were observed according to the main job activity: in pool attendants, THMs alveolar air were approximately double those observed in employees working in other areas of the swimming pools (25.1+/-16.5 microg/m3 vs. 14.8+/-12.3 microg/m3, P < 0.01). THMs in alveolar air samples were significantly correlated with THMs concentrations in ambient air (r = 0.57; P < 0.001). Indoor swimming pool employees are exposed to THMs at ambient air levels higher than the general population. The different environmental exposure inside the swimming pool can induce a different internal dose in exposed workers. The correlation found between ambient and alveolar air samples confirms that breath analysis is a good biological index of occupational exposure to these substances at low environmental levels.

Adult↗

Long-lasting delayed hyperalgesia after subchronic swim stress.

Rats subjected to an inescapable subchronic stress, consisting of 10-20 min of forced swimming for 3 days, showed a thermal hyperalgesia and an enhanced nociceptive behavior to the subcutaneous administration of formalin 24 and 48 h, respectively, after the last swim session. Hyperalgesia to thermal and chemical stimulants was still present 8 and 9 days after the last swim session, respectively. Chemical, but not thermal, nociception was negatively correlated with the swim effort or struggle times during the last swim session. The serotonin-selective reuptake inhibitors clomipramine (2.5 mg/kg/day, i.p., started 3 or 7 days before stress) and fluoxetine (0.25 mg/kg/day, i.p., started 7 days before stress), or serotonin precursor tryptophan (3 mg/kg/day, i.p., 24 h before each swim stress) blocked the development of both the thermal and the chemical hyperalgesia and increased swim effort times compared to vehicle-treated rats. These treatments did not affect nociceptive responses in control rats subjected to sham swimming. These findings suggest that repeated stress can produce a long-lasting increase in pain sensitivity to both phasic or tonic noxious stimuli by diminishing central serotonin activity. This model may help elucidate the underlying neural mechanisms that mediate the effects of repeated stress on pain sensitivity and affective states.

Animals↗