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PubMed1998

Analysis of the central pattern generator for swimming in the mollusk Clione.

The pteropod mollusk Clione limacina swims by rhythmic movements of two wings. The central pattern generator (CPG) for swimming, located in the pedal ganglia, is formed by three groups of interneurons. The interneurons of the groups 7 and 8 are of crucial importance for rhythm generation. They are endogenous oscillators capable of generating rhythmic activity with a range of frequencies typical of swimming after extraction from the ganglia. This endogenous rhythmic activity is enhanced by serotonin. The interneurons 7 and 8 produce one prolonged action potential (about 100 ms in duration) per cycle. Prolonged action potentials contribute to determining the duration of the cycle phases. The interneurons of two groups inhibit one another determining their reciprocal activity. The putative transmitters of groups 7 and 8 interneurons are glutamate and acetylcholine, respectively. Transition from one phase to the other is facilitated by the plateau interneurons of group 12 that contribute to termination of one phase and to initiation of the next phase. Maintaining the rhythm generation and transition from one phase to the other is also promoted by postinhibitory rebound. The redundant organization of the swimming generator guarantees the high reliability of its operation. Generation of the swimming output persisted after the inhibitory input from interneurons 8 to 7 had been blocked by atropine. Activity of the swimming generator is controlled by a set of command neurons that activate, inhibit or modulate the operation of the swimming CPG in relation to a behaviorally relevant context.

Animals↗
PubMed1984

Fictive locomotion in the lamprey spinal cord in vitro compared with swimming in the intact and spinal animal.

A comparison has been made of the patterns of muscle activity during swimming in the intact and spinal lamprey, and the patterns of ventral root activity in the in vitro preparation of the lamprey spinal cord. Electromyographic (e.m.g.) activity was recorded with intramuscular bipolar electrodes from three segmental levels in intact lampreys swimming in a swim-mill at a range of swimming speeds. The patterns of activity obtained were similar to those seen in elasmobranch and teleost fish. After high spinal transection, lampreys could be induced to swim continuously for a period of several minutes in the swim-mill by a light initial mechanical stimulation of the tail or dorsal fin. The patterns of e.m.g. activity obtained from spinal animals at a range of swimming speeds were similar to those obtained in the intact state. Portions of spinal cord were isolated encompassing those segments from which e.m.g. recordings had been made and ventral root recordings were made in vitro of the rhythmic activity induced by bath application of D-glutamate. In all experiments the mean duration of the bursts of activity at any segmental level was directly proportional to the mean cycle duration, and the constant of proportionality (about 0.36) was similar for all three types of preparation. In all preparations the mean time delay for the activation of segments in the rostral-caudal direction was proportional to the cycle duration and to the number of segments between recording positions. The proportionality constant, or phase lag per segment, was approximately equal to 0.01 in all three types of preparation.

Action Potentials↗
PubMed2005

Modification of cycling biomechanics during a swim-to-cycle trial.

The aim of this study was to investigate the effects of drafting, i.e., swimming directly behind a competitor, on biomechanical adaptation during subsequent cycling. Eight well-trained male triathletes underwent three submaximal sessions in a counterbalanced order. These sessions comprised a 10-min ride on a bicycle ergometer at 75% of maximal aerobic power (MAP) at a freely chosen cadence. This exercise was preceded either by a 750-m swim performed alone at competition pace (SCA trial; swimming-cycling alone), a 750-m swim in a drafting position at the same pace as during SCA (SCD trial; swimming-cycling with drafting), or a cycling warm-up at 30% of MAP for the same duration as the SCA trial (CTRL trial). The results indicated that the decrease in metabolic load when swimming in a drafting position (SCD trial) was associated with a significantly lower pedal rate and significantly higher mean and peak resultant torques when compared to the SCA trial, p < 0.05. These results could be partly explained by the lower relative intensity during swimming in the SCD trial when compared with the SCA trial, involving a delayed manifestation of fatigue in the muscles of the lower limbs at the onset of cycling.

Adult↗
PubMed2001

Hypothermia during sports swimming in water below 11 degrees C.

OBJECTIVES: To assess precautions needed to avoid dangerous hypothermia in endurance sports swims in water below 11 degrees C, using rectal temperature, anthropometric measurements, and voluntary swim times during a six day marathon relay swim. METHODS: The time in the water and the decrease in rectal temperature were measured during the longest of three to five relay swims by each of eight experienced swimmers in 9.4-11.0 degrees C water. Height, weight, and four skinfold thicknesses were measured. RESULTS: Swimmers with less subcutaneous fat terminated their swims after significantly less time in the water than those with thicker skinfold thickness, even though their rectal temperatures were not significantly lower. The lowest rectal temperature recorded was 34.3 degrees C. CONCLUSIONS: Subjective sensation in these experienced swimmers gave reliable guidance on safe durations for swims, and all voluntarily left the water with rectal temperatures that present no threat to people able to rewarm in safe surroundings. Endurance swims in highly competitive conditions or water below 9 degrees C may require continuous temperature monitoring for safety.

Adult↗
PubMed1969

Sudden failure of swimming in cold water.

To investigate the effect of cold water on swimming four men who declared themselves good swimmers were immersed fully clothed on separate days in water at 23.7 degrees and 4.7 degrees C. The time that they were able to swim in the cold water was much shorter than in the warm. The two shortest swims ended after 1.5 and 7.6 minutes, before rectal temperature fell, when the men suddenly floundered after developing respiratory distress with breathing rates of 56-60/min. The other cold swims, by the two fattest men, ended less abruptly with signs of general and peripheral hypothermia.It is concluded that swimming in cold water was stopped partly by respiratory reflexes in the thin men and hypothermia in the fat, and partly by the cold water's high viscosity. The longer swimming times of the fat men are attributed largely to their greater buoyancy enabling them to keep their heads above water during the early hyperventilation.The findings explain some reports of sudden death in cold water. It is clearly highly dangerous to attempt to swim short distances to shore without a life-jacket in water near 0 degrees C.

Adult↗
PubMed1995

Risk of otitis externa after swimming in recreational fresh water lakes containing Pseudomonas aeruginosa.

OBJECTIVE: To determine whether an outbreak of otitis externa was due to bathing in recreational fresh water lakes and to establish whether the outbreak was caused by Pseudomonas aeruginosa in the water. DESIGN: Matched case-control study. SETTING: The Achterhoek area, the Netherlands. SUBJECTS: 98 cases with otitis externa and 149 controls matched for age, sex, and place of residence. MAIN OUTCOME MEASURES: Odds ratios for type of swimming water and frequency of swimming; presence of P aeruginosa in ear swabs and fresh water lakes. RESULTS: Otitis externa was strongly associated with swimming in recreational fresh water lakes in the previous two weeks (odds ratio 15.5 (95% confidence interval) 4.9 to 49.2) compared with non-swimming). The risk increased with the number of days of swimming, and subjects with recurrent ear disease had a greatly increased risk. The lakes met the Dutch bathing water standards and those set by the European Commission for faecal pollution in the summer of 1994, but P aeruginosa was isolated from all of them, as well as from the ear swabs of 78 (83%) of the cases and 3 (4%) of the controls. CONCLUSIONS: Even when current bathing water standards are met, swimming can be associated with a substantial risk of otitis externa because of exposure to P aeruginosa. People with recurrent ear disease should take special care when swimming in waters containing P aeruginosa.

Adult↗
PubMed1999

Effects of swimming training on aerobic capacity and exercise induced bronchoconstriction in children with bronchial asthma.

BACKGROUND: A study was undertaken to determine whether swimming training improved aerobic capacity, exercise induced bronchoconstriction (EIB), and bronchial responsiveness to inhaled histamine in children with asthma. METHODS: Eight children with mild or moderate asthma participated in swimming training every day for six weeks. The intensity of training was individually determined and set at 125% of the child's lactate threshold (LT), measured using a swimming ergometer. Another group of eight asthmatic children served as control subjects. Aerobic capacity and the degree of EIB were assessed by both cycle ergometer and swimming ergometer before and after swimming training. RESULTS: The mean (SD) aerobic capacity at LT increased by 0.26 (0.11) kp after training when assessed with the swimming ergometer and by 10.6 (4.5) W when assessed with the cycle ergometer, and these changes were significantly different from the control group. The mean (SD) maximum % fall in forced expiratory volume in one second (FEV1) to an exercise challenge (cycle ergometer) set at 175% of LT decreased from 38.7 (15.4)% before training to 17.9 (17.6)% after training, but with no significant difference from the control group. There was, however, no difference in histamine responsiveness when compared before and after the training period. CONCLUSION: A six week swimming training programme has a beneficial effect on aerobic capacity but not on histamine responsiveness in children with asthma.

Asthma↗
PubMed2000

Upregulation of neuronal nitric oxide synthase in skeletal muscle by swim training.

Exercise enhances cardiac output and blood flow to working skeletal muscles but decreases visceral perfusion. The alterations in nitric oxide synthase (NOS) activity and/or expression of the cardiopulmonary, skeletal muscle, and visceral organs induced by swim training are unknown. In sedentary and swim-trained rats (60 min twice/day for 3-4 wk), we studied the alterations in NOS in different tissues along with hindquarter vasoreactivity in vivo during rest and mesenteric vascular bed reactivity in vitro. Hindquarter blood flow and conductance were reduced by norepinephrine in both groups to a similar degree, whereas N(G)-nitro-L-arginine methyl ester reduced both indexes to a greater extent in swim-trained rats. Vasodilator responses to ACh, but not bradykinin or S-nitroso-N-acetyl-penicillamine, were increased in swim-trained rats. Ca(2+)-dependent NOS activity was enhanced in the hindquarter skeletal muscle, lung, aorta, and atria of swim-trained rats together with increased expression of neuronal NOS in the hindquarter skeletal muscle and endothelial NOS in the cardiopulmonary organs. Mesenteric arterial bed vasoreactivity was unaltered by swim training. Physiological adaptations to swim training are characterized by enhanced hindquarter ACh-induced vasodilation with upregulation of neuronal NOS in skeletal muscle and endothelial NOS in the lung, atria, and aorta.

Animals↗
PubMed1978

Estimation of cardiac output by the CO2 rebreathing method during tethered swimming.

The reproducibility of cardiac output (Q) estimated by the CO2 rebreathing method during tethered swimming was studied in five highly trained college swimmers. The reproducibility of the CO2 rebreathing method for estimations of Q during tethered swimming was similar to the reproducibility reported for the CO2 rebreathing method, direct Fick method, or dye-dilution method during either cycling or treadmill walking. All duplicate estimates of Q by the CO2 rebreathing method were within 15% of one another. A comparison was made between the Q's estimated by the CO2 rebreathing method during tethered swimming and previously published data on Q determined by the dye-dilution method during free swimming in a flune. At any given oxygen uptake, Q obtained by the CO2 rebreathing method during tethered swimming was not significantly different from the Q obtained by the dye-dilution method during flume swimming. Estimates of Q by the CO2 rebreathing method made during high intensities of tethered swimming were reproducible and appear to be valid.

Adult↗
PubMed1987

Patterns of EMG activity of rat plantaris muscle during swimming and other locomotor activities.

The purpose of the study was to examine the patterns of electromyographic (EMG) activity of the rat plantaris during loaded swimming in comparison with other locomotor activities. Five female Sprague-Dawley rats were implanted with chronic bipolar electrodes in the plantaris muscle of the left hindlimb under pentobarbital anesthesia. Characteristics of EMG bursts recorded while the conscious rat was performing treadmill walking (0.24 m/s) were stable and reproducible 10-14 days postsurgery. Following this stabilization period, records of EMG activity were obtained during walking, loaded swimming (6.5 g attached to tail), and several other locomotor tasks. Compared to walking, EMG bursts during loaded swimming were significantly higher (67%) in maximum amplitude, one-third as long in duration, and occurred at a greater rate (4.4 vs. 1.7 bursts/s, P less than 0.05). Swimming bursts were of higher amplitudes than those of all other activities examined and reached 65% of the EMG amplitude recorded following stimulation of the sciatic nerve with supramaximal voltage. The addition of a mass to the animal's tail during swimming did not increase the EMG burst amplitudes but resulted in a higher frequency of bursts. Compared with treadmill walking, loaded swimming elicited burst of high variability in amplitude. Swimming in the rat involves rapid, extensive activation of plantaris, thus providing an exercise model to study the adaptability of the neuromuscular system to prolonged activity of this type.

Animals↗
PubMed2002

Declines in physiological functional capacity with age: a longitudinal study in peak swimming performance.

We followed up swimming performance times of 321 women and 319 men who participated in the US Masters Swimming Championships over a 12-yr period. All swimmers placed in the top 10 in their age group over 3 yr (mean = 5 yr). A random coefficients model for repeated measures was used to derive a line of best fit from a group of regression lines for each subject. Both 50- and 1,500-m swimming performance declined modestly until approximately 70 yr of age, where a more rapid decline was observed in both men and women. Compared with 1,500-m swimming, the 50-m freestyle declined more modestly and slowly with age. The rate and magnitude of declines in swimming performance with age were greater in women than in men in 50-m freestyle; such sex-related differences were not observed in 1,500-m freestyle. Overall, the variability along a population regression line increased markedly with advancing age. The present longitudinal findings indicate that 1) swimming performance declines progressively until age 70, where the decrease becomes quadratic; 2) the rates of the decline in swimming performance with age are greater in a long-duration than in a short-duration event, suggesting a relatively smaller loss of anaerobic muscular power with age compared with cardiovascular endurance; 3) the age-related rates of decline are greater in women than in men only in a short-duration event; and 4) the variability of the age-related decline in performance increases markedly with advancing age.

Adult↗
PubMed2002

Activation of embryonic red and white muscle fibers during fictive swimming in the developing zebrafish.

Sub-threshold, motoneuron-evoked synaptic activity was observed in zebrafish embryonic red (ER) and white (EW) muscle fibers paralyzed with a dose of D-tubocurarine insufficient to abolish synaptic activity to determine whether muscle activation was coordinated to produce the undulating body movements required for locomotion. Paired whole-cell recordings revealed a synaptic drive that alternated between ipsilateral and contralateral myotomes and exhibited a rostral-caudal delay in timing appropriate for swimming. Both ER and EW muscle were activated during fictive swimming. However, at the fastest fictive swimming rates, ER fibers were de-recruited, whereas they could be active in isolation of EW fibers at the slowest fictive swimming rates. Prior to hatching, fictive swimming was preceded by a lower frequency, more robust and rhythmic synaptic drive resembling the "coiling" behavior of fish embryos. The motor activity observed in paralyzed zebrafish closely resembled the swimming and coiling behaviors observed in these developing fishes. At the early developmental stages examined in this study, myotomal muscle recruitment and coordination were similar to that observed in adult fishes during swimming. Our results indicate that the patterned activation of myotomal muscle is set from the onset of development.

Animals↗
PubMed1983

Swimming rhythm in decerebrated, paralyzed stingrays: normal and abnormal coupling.

Rhythmic motoneuronal activity was recorded from decerebrated, paralyzed stingrays and compared with electromyograms recorded from the same animals. Before and after paralysis, a rostral-to-caudal sequence of alternation occurred between dorsal (elevator) and ventral (depressor) efferents. The swimming pattern was thus observed in the absence of phasic afferent input, and this constitutes fictive locomotion. After paralysis, both the intersegmental delay (time between activation at progressively caudal recording sites) and the burst duration remained linearly related to the swim cycle period. In many instances, neither the slope nor the intercept was significantly altered by immobilization. The intercepts all fell near the origin, indicating that the fictive rhythm remains constant phase coupled. Although the swimming rhythm was obtained after paralysis, some differences occurred. These included fewer and shorter spontaneous sequences, a greater range of cycle periods, and longer burst durations. During fictive swimming, the burst duration:cycle period ratio usually increased to 0.53 from 0.39 observed before paralysis. Therefore, the silent periods seen between burst discharges in antagonist efferents during movement were often absent after paralysis. Mechanical stimulation of the tail reduced both cycle periods and burst durations; however, the burst:cycle ratio remained greater than or equal to 0.50. The linear relation between burst duration and cycle period found for spontaneous sequences was not changed by stimulation of the tail. During fictive swimming the inter- and intrasegmental coupling that characterizes stingray swimming becomes labile. Abnormal coupling appears more often during sequences with long swim cycles. Intrasegmental coupling is tighter than intersegmental coupling at any cycle period. Rhythmic activity at one segmental level can be independent of activity at other levels. This suggests that multiple oscillator circuits exist that are not dependent on propriospinal circuits interconnecting different segments. Rhythmicity in elevator and depressor motoneurons is not dependent on reciprocal connections between the circuitry driving the motor nuclei. Therefore, separate oscillators for elevators and depressors appear to be present within one spinal segment.

Animals↗
PubMed1984

Spinal cord pathways involved in initiation of swimming in the stingray, Dasyatis sabina: spinal cord stimulation and lesions.

In spinally transected stingrays, electrical stimulation of a site just ventral to the dorsal root entry zone or a site in the intermediate portions of the lateral funiculus produced rhythmic swimming like movements of the contralateral pectoral fin. Electromyographic (EMG) records collected during cord-stimulated rhythms had the same pattern of activity and sometimes the same intersegmental coordination as those collected during spontaneous swimming of the same animal. In paralyzed, high-spinal stingrays, the only stimulation sites that produced rhythmic activity (fictive swimming) in the pectoral fin motor nerves were in the intermediate portion of the lateral funiculus. The evoked rhythm occurred in the motor nerves that were contralateral to the stimulated side of the spinal cord. The effects of subtotal lesions of the rostral spinal cord on spontaneous swimming behavior were assessed by analysis of EMG records taken before and after the lesions were made. Severe deficits in swimming occurred after bilateral ablation of intermediate portions of the lateral funiculi. In agreement with previous results, the stimulation experiments indicate that the stingray spinal cord contains an inherent capacity to generate properly coordinated rhythmic swimming. The current experiments also suggest that the descending pathways(s) that normally functions to initiate swimming projects through the intermediate aspects of the lateral funiculi.

Animals↗
PubMed1996

Local effects of glycinergic inhibition in the spinal cord motor systems for swimming in amphibian embryos.

1. We have studied the effects of locally applying the glycinergic antagonist strychnine to rhythmically active spinal neurons in amphibian embryos during fictive swimming. Intracellular recordings were made from motoneurons and premotor interneurons in Xenopus laevis, a well-studied model system, and from motoneurons in three other species (Rana temporaria, Bufo bufo, and Triturus vulgaris). Overall, these embryos cover a range of swimming patterns from the short-cycle-period, brief-motor-root bursts of Xenopus, to the long-cycle-period, long-motor-root bursts of Rana, which are more typical of adult patterns. 2. Local strychnine application had no significant effect on the gross pattern of swimming; episode duration and the burst duration in rostral ventral roots away from the application site were unaltered, and left-right alternation was preserved. We have therefore been able to examine the effects of inhibition on individual neurons, uninfluenced by overall changes in the operation of the swimming neural circuitry. 3. In all cases strychnine blocked midcycle inhibition and significantly increased the peak on-cycle depolarization during swimming. In Rana, Bufo, and Triturus motoneurons, and in Xenopus interneurons, strychnine significantly increased the reliability of firing during swimming. In Xenopus motoneurons, where spiking was 100% reliable anyway, the timing of the spikes was advanced relative to rostral ventral root activity. These results do not provide support for postinhibitory rebound as a factor in the spike-generating process during swimming. In addition to midcycle inhibition, Xenopus motoneurons can also show a smaller, additional on-cycle inhibition that is blocked by strychnine. 4. In both Rana and Bufo the duration of caudal ventral root bursts close to the site of drug application was increased by strychnine, showing that the increased motoneuron reliability not only leads to more intense, but also more extensive, ventral root activity. 5. At the level of single neurons, glycinergic inhibition effectively reduces on-cycle excitation and in turn controls the reliability, extent, and precise timing of motoneuron firing. These changes may be the individual components underlying broader effects of inhibition described previously, such as locomotor frequency control. They also show how any modulation of inhibition in localized regions of the spinal cord could produce localized control of neuronal firing properties.

Amphibians↗
PubMed1997

Spinal cord coordination of hindlimb movements in the turtle: intralimb temporal relationships during scratching and swimming.

Spinal cord neuronal circuits generate motor neuron activity patterns responsible for rhythmic hindlimb behaviors such as scratching and swimming. Kinematic analyses of limb movements generated by this motor neuron output reveal important characteristics of these behaviors. Intralimb kinematics of the turtle hindlimb were characterized during five distinct rhythmic forms of behavior: three forms of scratching and two forms of swimming. In each movement cycle for each form, the angles of the hip and knee joints were measured as well as the timing of a behavioral event, e.g., rub onset in scratching or powerstroke onset in swimming. There were distinct differences between the kinematics of different forms of the same behavior, e.g., rostral scratch versus pocket scratch. In contrast, there were striking similarities between forms of different behaviors, e.g., rostral scratch versus forward swimming. For each form of behavior there was a characteristic angular position of the hip at the onset of each behavioral event (rub or powerstroke). The phase of the onset of knee extension within the hip position cycle occurred while the hip was flexing in the rostral scratch and forward swim and while the hip was extending in the pocket scratch, caudal scratch, and back-paddling form of swimming. The phase of the onset of the behavioral event was not statistically different between rostral scratch and forward swim; nor was it different between pocket scratch and caudal scratch. These observations of similarities at the movement level support the suggestion that further similarities, such as shared spinal circuitry, may be present at the neural circuitry level as well.

Animals↗
PubMed1999

Segmental distribution of common synaptic inputs to spinal motoneurons during fictive swimming in the lamprey.

These experiments were designed to measure the degree of shared synaptic inputs coming to pairs of myotomal motoneurons during swimming activity in the isolated spinal cord of the lamprey. In addition, the experiments measured the decrease in the degree of shared synaptic inputs with the distance between the motoneurons to assess the segmental distribution of these shared inputs. Intracellular microelectrode recordings of membrane potential were made simultaneously on pairs of myotomal motoneurons during swimming activity induced with an excitatory amino acid. The swim cycle oscillations of motoneuron membrane potentials were removed with a digital notch filter, thus leaving the fast synaptic activities that underlie these slower oscillations. Cross-correlations of the fast synaptic activities in two simultaneously recorded motoneurons were made to measure the degree of shared inputs. The cross-correlation was done on time windows restricted to one swim cycle or to part of a swim cycle, and 50 consecutive swim cycle cross-correlograms then were averaged. The peak coefficients of the cross-correlations exhibited a wide range, even for pairs of motoneurons located near one another (range = 0.06-0.74, for pairs located within 2 segments). This observation suggests that there may be different functional classes of myotomal motoneurons with inputs originating from different sets of premotor interneurons. In spite of this variability, the mean peak correlation coefficients of motoneuron pairs clearly decreased with the distance between them. With separations of more than five segments, there was little or no clear correlation between the motoneurons (range = 0.04-0.10). These results suggest that common synaptic inputs to motoneurons during fictive swimming originate from local premotor interneurons and that beyond five spinal segments, common premotor inputs are rare or weak to motoneurons. Thus the premotor signals originating from the locomotor network have relatively short distribution lengths, on the order of 5 segments of 120 total spinal segments.

Animals↗
PubMed2000

Escape swim network interneurons have diverse roles in behavioral switching and putative arousal in Pleurobranchaea.

Escape swimming in the predatory sea slug Pleurobranchaea is a dominant behavior that overrides feeding, a behavioral switch caused by swim-induced inhibition of feeding command neurons. We have now found distinct roles for the different swim interneurons in acute suppression of feeding during the swim and in a longer-term stimulation of excitability in the feeding network. The identified pattern-generating swim neurons A1, A3, A10, and their follower interneuron A-ci1, suppress feeding motor output partly by excitation of the I1 feeding interneurons, which monosynaptically inhibit both the feeding command neurons, PC(P), PSE, and other major interneurons, the I2s. This mechanism exerts broad inhibition of the feeding network suitable to an escape response; broader than feeding suppression in learned and satiation-induced food avoidance and acting through a different presynaptic pathway. Four intrinsic neuromodulatory neurons of the swim network, the serotonergic As1-4, add little to direct suppression of feeding. Rather, they monosynaptically excite the serotonergic metacerebral giant (MCG) neurons of the feeding network, themselves intrinsic neuromodulators of feeding, as well as a cluster of adjacent serotonergic feeding neurons, with both fast and slow EPSPs. They also provide mild neuromodulatory excitation of the PC(P)/PSE feeding command neurons, and I1 and I2 feeding interneurons, which is masked by inhibition during the swim. As1-4 also excite the serotonergic pedal ganglion G neurons for creeping locomotion. These observations further delineate the nature of the putative serotonergic arousal system of gastropods and suggest a central coordinating role to As1-4.

Animals↗