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Central pattern generator for swimming in Melibe.

The nudibranch mollusc Melibe leonina swims by bending from side to side. We have identified a network of neurons that appears to constitute the central pattern generator (CPG) for this locomotor behavior, one of only a few such networks to be described in cellular detail. The network consists of two pairs of interneurons, termed 'swim interneuron 1' (sint1) and 'swim interneuron 2' (sint2), arranged around a plane of bilateral symmetry. Interneurons on one side of the brain, which includes the paired cerebral, pleural and pedal ganglia, coordinate bending movements toward the same side and communicate via non-rectifying electrical synapses. Interneurons on opposite sides of the brain coordinate antagonistic movements and communicate over mutually inhibitory synaptic pathways. Several criteria were used to identify members of the swim CPG, the most important being the ability to shift the phase of swimming behavior in a quantitative fashion by briefly altering the firing pattern of an individual neuron. Strong depolarization of any of the interneurons produces an ipsilateral swimming movement during which the several components of the motor act occur in sequence. Strong hyperpolarization causes swimming to stop and leaves the animal contracted to the opposite side for the duration of the hyperpolarization. The four swim interneurons make appropriate synaptic connections with motoneurons, exciting synergists and inhibiting antagonists. Finally, these are the only neurons that were found to have this set of properties in spite of concerted efforts to sample widely in the Melibe CNS. This led us to conclude that these four cells constitute the CPG for swimming. While sint1 and sint2 work together during swimming, they play different roles in the generation of other behaviors. Sint1 is normally silent when the animal is crawling on a surface but it depolarizes and begins to fire in strong bursts once the foot is dislodged and the animal begins to swim. Sint2 also fires in bursts during swimming, but it is not silent in non-swimming animals. Instead activity in sint2 is correlated with turning movements as the animal crawls on a surface. This suggests that the Melibe motor system is organized in a hierarchy and that the alternating movements characteristic of swimming emerge when activity in sint1 and sint2 is bound together.

Animals↗

Swimming behavior of the nudibranch Melibe leonina.

Swimming in the nudibranch Melibe leonina consists of five types of movements that occur in the following sequence: (1) withdrawal, (2) lateral flattening, (3) a series of lateral flexions, (4) unrolling and swinging, and (5) termination. Melibe swims spontaneously, as well as in response to different types of aversive stimuli. In this study, swimming was elicited by contact with the tube feet of the predatory sea star Pycnopodia helianthoides, pinching with forceps, or application of a 1 M KCl solution. During an episode of swimming, the duration of swim cycles (2.7 +/- 0.2 s [mean +/- SEM], n = 29) and the amplitude of lateral flexions remained relatively constant. However, the latency between the application of a stimulus and initiation of swimming was more variable, as was the duration of an episode of swimming. For example, when touched with a single tube foot from a sea star (n = 32), the latency to swim was 7.0 +/- 2.4 s, and swimming continued for 53.7 +/- 9.4 s, whereas application of KCl resulted in a longer latency to swim (22.3 +/- 4.5 s) and more prolonged swimming episodes (174.9 +/- 32.1 s). Swimming individuals tended to move in a direction perpendicular to the long axis of the foot, which propelled them laterally when they were oriented with the oral hood toward the surface of the water. The results of this study indicate that swimming in Melibe, like that in several other molluscs, is a stereotyped fixed action pattern that can be reliably elicited in the laboratory. These characteristics, along with the large identifiable neurons typical of many molluscs, make swimming in this nudibranch amenable to neuroethological analyses.

Animals↗

Effects of physical training and mental practice of in-clothes swimming: assessment by physiological parameters.

PURPOSE: This study examined whether a short-term physical or mental training reduces the physiological load and perceived exertion of the in-clothes swimming. METHODS: The study included 24 male inter-collegiate competitive swimmers with no previous experience of the in-clothes swimming. Prior to the training, the subjects performed 200-meter swimming with two styles of swimming, namely the crawl and elementary backstroke, and the degree of perceived exertion in the Borg scale, heart rate, and blood lactic acid level were determined. Following this baseline determination, the subjects were divided into 4 groups with 6 individuals each. These four groups were Group A "in-clothes training", Group B "image training", Group C "swimsuit training", and Group N "no training". Group A and C were lectured on the in-clothes swimming and practiced 7.5 min-long in-clothes swimming per day for a week with the two swimming styles, with Group A subjects wearing daily clothes and Group C subjects in swimsuits. Group B received 15 min of nonphysical mental practice primarily through viewing video recording of swimming performance everyday for a week. No training was given to Group N. At the completion of the training session, the subjects underwent the 2nd 200-meter swimming, and the physiological parameters were determined. RESULTS: The 1st in-clothes swimming load test showed that the elementary backstroke swimming resulted in significantly lower values of heart rate, blood lactic acid level, and perceived exertion than the crawl. For Group N, no difference was observed in the physiological parameters between the 1st and 2nd load test with either the elementary backstroke or crawl. Upon the 2nd in-clothes load test with the elementary backstroke, all three parameters were lower for Group A, B, and C than those seen for the 1st load test, and these differences were statistically significant, except for blood lactic acid in Group A. The 2nd in-clothes load test with the crawl showed that both heart rate and blood lactic acid were lower than those of the 1st load test for all three groups, with the differences in heart rate and blood lactic acid in Group A and that in blood lactic acid in Group B reaching the statistic significance. With respect to perceived exertion, the Borg score determined after the in-clothes load test with the crawl was significantly reduced for Group A, B, and C. The score after the in-clothes load test with the elementary backstroke was significantly reduced only for Group A. CONCLUSION: The elementary backstroke, when practiced in-clothes, imposes a relatively less physiological load. The present training methods can reduce physiological load and/or perceived exertion of the in-clothes swimming. Thus, both image training and swimsuit swimming are equally effective as a training method of the in-clothes swimming.

Adult↗

Interaction between the forced swimming test and fluoxetine treatment on extracellular 5-hydroxytryptamine and 5-hydroxyindoleacetic acid in the rat.

We used in vivo microdialysis to examine extracellular levels of 5-hydroxytryptamine (5-HT) and 5-hydroxyindoleacetic acid (5-HIAA) in the striatum and the lateral septum during the forced swimming test, (FST) a behavioral test conducted in rats that is commonly used to predict the effect of antidepressant drugs. The forced swimming test consisted of a 15-min pretest swim and a 5-min test swim 24 hr later. The antidepressant fluoxetine (20 mg/kg s.c.) or saline was administered 23.5, 5 and 1 hr before the test swim. In the striatum, the pretest swim increased 5-HT in both treatment groups. On the second day, the test swim had no effect on 5-HT in saline-treated rats but slightly decreased striatal 5-HT in fluoxetine-treated rats. In the lateral septum, the pretest swim decreased 5-HT in both treatment groups. On the second day, the test swim had no effect on 5-HT in saline-treated rats but decreased lateral septum 5-HT in fluoxetine-treated rats. Ratings of behavior showed that fluoxetine treatment increased swimming behavior and decreased immobility during the test swim. Immobility was positively correlated and swimming was negatively correlated with changes in extracellular 5-HT in the lateral septum but not in the striatum. Therefore, fluoxetine treatment altered adaptation of the regional response of extracellular 5-HT ordinarily produced in the FST, reversing the 5-HT response to the initial swim in the striatum and restoring the response to the initial swim in the lateral septum.

Animals↗

A respirometric analysis of fuel use during aerobic swimming at different temperatures in rainbow trout (Oncorhynchus mykiss)

Instantaneous fuel usage at 5 degreesC or 15 degreesC was assessed by measurement of rates of O2 consumption (O2), CO2 excretion (CO2) and nitrogenous waste excretion (nitrogen =ammonia-N + urea-N) in juvenile rainbow trout (Oncorhynchus mykiss) at rest and during swimming at 45 % and 75 % of aerobic capacity (Ucrit). After 2 weeks of training at approximately 1 body length s-1 (BL s-1), critical swimming speeds (approximately 3.0 BL s-1) and whole-body energy stores (total protein, lipids and carbohydrates) were identical in fish acclimated to 5 degreesC or 15 degreesC. O2 and CO2 increased with swimming speed at both temperatures and were higher at 15 degreesC than at 5 degreesC at all speeds, but the overall Q10 values (1.23-1.48) were low in these long-term (6 weeks) acclimated fish. The respiratory quotient (CO2/O2, approximately 0.85) was independent of both temperature and swimming speed. In contrast to O2 and CO2, the rate of ammonia excretion was independent of swimming speed, but more strongly influenced by temperature (Q10 1. 4-2.8). Urea excretion accounted for 15-20 % of nitrogen, was unaffected by swimming speed and showed a tendency (P<0.07) to be positively influenced by temperature at one speed only (45 % Ucrit). Nitrogen quotients (NQ nitrogen/O2) were generally higher in warm-acclimated fish, remaining independent of swimming speed at 15 degreesC (0.08), but decreased from about 0.08 at rest to 0.04 during swimming at 5 degreesC. Instantaneous aerobic fuel use calculations based on standard respirometric theory showed that both acclimation temperature and swimming speed markedly influenced the relative and absolute use of carbohydrates, lipids and proteins by trout. At rest, cold-acclimated trout used similar proportions of carbohydrates and lipids and only 27 % protein. During swimming, protein use decreased to 15 % at both speeds while the relative contributions of both lipid and carbohydrate increased (to more than 40 %). On an absolute basis, carbohydrate was the most important fuel for fish swimming at 5 degreesC. In contrast, resting fish acclimated to 15 degreesC utilized 55 % lipid, 30 % protein and only 15 % carbohydrate. However, as swimming speed increased, the relative contribution of carbohydrate increased to 25 %, while the protein contribution remained unchanged at approximately 30 %, and lipid use decreased slightly (to 45 %). On an absolute basis, lipid remained the most important fuel in fish swimming at 15 degreesC. These results support the concept that lipids are a major fuel of aerobic exercise in fish, but demonstrate that the contribution of protein oxidation is much smaller than commonly believed, while that of carbohydrate oxidation is much larger, especially at higher swimming speeds and colder temperature.

Journal Article↗

Reboxetine attenuates forced swim test-induced behavioural and neurochemical alterations in the rat.

The forced swim test is a behavioural paradigm that is predicative of antidepressant activity in rodents. Until recently, research has focused on the ability of antidepressant drugs to decrease immobility in the forced swim test paradigm, but the neurochemical sequelae induced by swim stress, or the neurochemical basis of antidepressant-induced behavioural changes have received little attention. In this regard, we have recently demonstrated that forced swim test exposure increases serotonergic activity in the amygdala, frontal cortex and hippocampus and dopamine turnover in the striatum. In addition, forced swim test-exposure activates the hypothalamic pituitary adrenal axis. The purpose of the present study was to examine the effect of treatment with the selective noradrenaline reuptake inhibitor reboxetine (3, 10 and 30 mg/kg; i.p.) on immobility and defaecation scores in the forced swim test, and on forced swim test-induced neurochemical and hypothalamic pituitary adrenal axis changes in the rat. Reboxetine treatment (10 and 30 mg/kg) significantly decreased immobility and defaecation in the forced swim test in dose dependent manner. Furthermore, reboxetine produced a dose dependent attenuation of forced swim test-induced increases in serotonin turnover in the amygdala and frontal cortex and dopamine turnover in the striatum. Reboxetine (30 mg/kg) produced a modest, but non-significant, attenuation of forced swim test-induced increases in serum corticosterone concentrations. These data demonstrate that, in addition to the behavioural activity of reboxetine in the rat forced swim test paradigm, a dose-dependent attenuation of swim stress-induced increases in serotonergic and dopaminergic activity occurred in a region specific manner. These are the first data to demonstrate that treatment with the selective noradrenaline reuptake inhibitor, reboxetine can impact on the activity of other neurotransmitter systems in response to stress. Moreover, these data further demonstrate that this paradigm is a valuable tool in studying the effect of antidepressants, on both behaviour and swim stress-related alterations in central neurotransmitter function and hypothalamic pituitary adrenal axis activity in the rat.

3,4-Dihydroxyphenylacetic Acid↗

Simulated front crawl swimming performance related to critical speed and critical power.

PURPOSE: Competitive pool swimming events range in distance from 50 to 1500 m. Given the difference in performance times (+/- 23-1000 s), the contribution of the aerobic and anaerobic energy systems changes considerably with race distance. In training practice the regression line between swimming distance and time (Distance = critical velocity x time + anaerobic swimming capacity) is used to determine the individual capacity of the aerobic and anaerobic metabolic pathways. Although there is confidence that critical velocity and anaerobic swimming capacity are fitness measures that separate aerobic and anaerobic components, a firm theoretical basis for the interpretation of these results does not exist. The purpose of this study was to evaluate the critical power concept and anaerobic swimming capacity as measures of the aerobic and anaerobic capacity using a modeling approach. METHODS: A systems model was developed that relates the mechanics and energetics involved in front crawl swimming performance. From actual swimming flume measurements, the time dependent aerobic and anaerobic energy release was modeled. Data derived from the literature were used to relate the energy cost of front crawl swimming to swimming velocity. A balance should exist between the energy cost to swim a distance in a certain time and the concomitant aerobic and anaerobic energy release. The ensuing model was used to predict performance times over a range of distances (50-1500 m) and to calculate the regression line between swimming distance and time. RESULTS AND CONCLUSIONS: Using a sensitivity analysis, it was demonstrated that the critical velocity is indicative for the capacity of the aerobic energy system. Estimates of the anaerobic swimming capacity, however, were influenced by variations in both anaerobic and aerobic energy release. Therefore, it was concluded that the anaerobic swimming capacity does not provide a reliable estimate of the anaerobic capacity.

Adolescent↗

Modulation of swimming activity in the medicinal leech by serotonin and octopamine.

1. The monoamines serotonin (5-HT) and octopamine (OA) enhance the expression of swimming activity in the medicinal leech (Willard, 1981; Belanger and Orchard, 1988). We explored further the effects of these monoamines and related agents on swimming activity observed in isolated leech nerve cords. 2. We confirmed that swimming activity is induced reversibly following exposure of the nerve cord to 5-HT (50 microM); the half-maximal rate of swimming activity develops in about 15 min. Swimming activity returns to control levels about 30 min after drug washout. 3. Swim-induction by 5-HT is blocked by the presence of 10 microM cyproheptadine (a 5-HT antagonist). 4. Although apparently less effective than 5-HT, OA application to nerve cords also induced swimming activity. 5. Depletion of endogenous amines from nerve cords by acute exposure to reserpine (10-150 microM) blocked stimulus-evoked swimming activity within 4 hr. 6. Subsequent application of 5-HT (50 microM) or OA (100 microM) reinstated stimulus-evoked swimming and induced repeated episodes of non-triggered swimming activity. 7. Application of cAMP and cAMP analogs, as well as phosphodiesterase inhibitors (theophylline and IBMX), mimicked the effects of the monoamines, suggesting that 5-HT and OA may activate swimming activity by increasing neuronal cAMP. 8. We obtained episodes of swim-like activity from individual, isolated ganglia exposed to 5-HT or OA. Such episodes were usually brief, with variable cycle period. 9. We conclude that individual nerve cord ganglia contain the complete neuronal circuitry required to generate the rudiments of swimming activity.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Sperm swim-up techniques and DNA fragmentation.

BACKGROUND: Swim-up techniques for sperm separation may have detrimental effects on sperm DNA. We wished to determine whether the normal swim-up method with centrifugation used in our laboratory, which involves a centrifugation step, was harmful to sperm compared with swim-up without centrifugation. METHODS: Semen samples were obtained from patients undergoing IVF or andrology assessment. An aliquot was removed for fixation and subsequent DNA fragmentation determination. The remaining sample was divided into two equal parts, which were subjected to swim-up either with (normal swim-up) or without (direct-swim-up) centrifugation. Semen analysis was performed both before and after swim-up. DNA fragmentation, in spermatozoa previously fixed in 4% paraformaldehyde, was assessed by the terminal transferase-mediated DNA end-labelling procedure (TUNEL). The percentage of spermatozoa with DNA damage after each swim-up technique was compared with that in the original semen sample. RESULTS: DNA damage was <5% in most samples. No significant change in DNA fragmentation was observed between the two swim-up procedures, although the 'normal' swim-up sample had significantly less DNA fragmentation than the pre-swim-up sample. CONCLUSIONS: We conclude that our normal swim-up technique caused no more DNA damage to spermatozoa from normal semen samples than a direct swim-up technique that involved no centrifugation step.

Adult↗

Effects of NMDA antagonists on memory processes in a novel two-trial swimming test.

Rats were placed in a circular swimming pool for two 3-min swims separated by 3 days. In the first swim control rats swam initially around the perimeter of the pool and later spent more time in the central region and swam more slowly. The time spent in the centre during min 1 was much higher in the second swim than in the first. This alteration of behaviour by the previous experience suggests that a memory of the first swim was formed. The NMDA antagonists, CPP (10-20mg/kg) or MK-801 (50-100µg/kg), administered before the first swim attenuated or reversed the decline in swimming speed and attenuated the intra-trial increase in the time spent in the centre. The alteration of swimming pattern between the two swims was also reduced. Administering the antagonists immediately after the first swim did not have this effect. Drug administration shortly before the second swim prevented the within-trial decline of swimming speed, but did not significantly reduce the high proportion of time spent in the central region during min 1. The results suggest that in this paradigm NMDA receptors are involved in within-trial habituation of swimming activity and in the initial stages of long-term memory formation, but are not involved in memory consolidation or retrieval.

Journal Article↗

[Exposure to chloroform in persons frequenting an indoor swimming pool].

Exposure to chloroform (CHCl3) in indoor swimming pool swimmers. CHCl3 presence has been evidenced many times in water and air of indoor pools as a consequence of water chlorination with sodium hypochlorite or other chlorinated disinfectants. In previous studies CHCl3 in environmental air appeared at a high concentration within one meter over the surface of the water and was correlated to the number of swimmers inside the pool. CHCl3 has also been evidenced in blood serum samples of people present at the swimming pool, at different levels depending on: the concentration in the environmental air, the intensity and the length of the sport activity and the number of swimmers in the pool. At the end of the swimming session CHCl3 concentration tends to lower quickly and usually is no longer detectable within half an hour. Other studies have evidenced CHCl3 in breath samples (alveolar air) of a volunteer subject who swam one hour every day in an indoor pool in Modena. The spirometric parameters of the same subject were evaluated in order to estimate his CHCl3 uptake during time spent at the swimming pool, both resting and swimming in different modes. In the present paper the A.A. report data about CHCl3 exposure, intake and uptake bin five agonistic swimmers (three males and two females), members of the same sport association, and regularly attending the same indoor swimming pool in Modena. The five subjects have been examined during four different sessions. Within every session the samples were collected in five different moments according to the following scheme: 1) Hygiene Institute: samples of blood, alveolar air, environmental air. 2) Swimming pool, after 1 hour sitting near the pool edge: samples of alveolar air, environmental air (within 1 m on the water surface). 3) Swimming pool, after 1 hour swimming: samples of blood, alveolar air, environmental air (within 1 m on the water surface). 4) Hygiene Institute, 1 hour after the end of the swimming time: samples of alveolar air, environmental air. 5) Hygiene Institute, 30 min after the previous sampling: samples of alveolar air, environmental air. Spirometric parameters have been evaluated in every subject, after resting and swimming (point 2 and 3). CHCl3 uptake has been calculated using the following formula: U = V(CI--CA)t where: U = uptake (microgram/h) V = pulmonary ventilation (l/min) CI = inspired (environmental) concentration (microgram/m3) CA = alveolar concentration (microgram/m3) t = exposure time (min) CHCl3 levels after swimming always appeared quite high; however low levels were evidenced also before exposure (point 1).(ABSTRACT TRUNCATED AT 400 WORDS)

Adolescent↗

Swimming by sea otters: adaptations for low energetic cost locomotion.

The energetics and hydrodynamics of surface and submerged swimming were compared in the sea otter (Enhydra lutris). 1. Sea otters used two distinct speed ranges that varied with swimming mode. Sustained surface swimming was limited to speeds less than 0.80 m/s, while sustained submerged swimming occurred over the range of 0.60 to 1.39 m/s. 2. Rates of oxygen consumption (VO2) at the transition speed (0.80 m/s) were 41% lower for submerged swimming by sea otters in comparison to surface swimming. 3. Total cost of transport for surface swimming sea otters, 12.56 joules/kg.m, was more than 12 times the predicted value for a similarly-sized salmonid fish. Transport costs for submerged swimming at the same speed was only 7.33 times the predicted value. 4. The allometric relationship for minimum cost of transport in surface swimming birds and mammals was y = 23.87 chi -0.15 where y = cost of transport in joules/kg.m and x = body mass in kg. This regression loosely parallels the relationship for salmonid fish. 5. Correlations between aquatic behavior, morphological specialization, and swimming energetics indicate that the development of swimming in mustelids involved transitions from fore-paw to hind-paw propulsion, and from surface to submerged swimming.

Animals↗

Stenotherms at sub-zero temperatures: thermal dependence of swimming performance in Antarctic fish.

We examined the burst swimming performance of two Antarctic fishes, Trematomus bernacchii and T. centronotus, at five temperatures between -1 degrees C and 10 degrees C. As Antarctic fishes are considered one of the most cold specialised and stenothermal of all ectotherms, we predicted they would possess a narrow thermal performance breadth for burst swimming and a correlative decrease in performance at high temperatures. Burst swimming was assessed by videotaping swimming sequences with a 50-Hz video camera and analysing the sequences frame-by-frame to determine maximum velocity, the distance moved throughout the initial 200 ms, and the time taken to reach maximum velocity. In contrast to our prediction, we found both species possessed a wide thermal performance breadth for burst swimming. Although maximum swimming velocity for both T. bernacchii and T. centronotus was significantly highest at 6 degrees C, maximum velocity at all other test temperatures was less than 20% lower. Thus, it appears that specialisation to a highly stable and cold environment is not necessarily associated with a narrow thermal performance breadth for burst swimming in Antarctic fish. We also examined the ability of the Antarctic fish Pagothenia borchgrevinki to acclimate their burst-swimming performance to different temperatures. We exposed P. borchgrevinki to either -1 degrees C or 4 degrees C for 4 weeks and tested their burst-swimming performance at four temperatures between -1 degrees C and 10 degrees C. Burst-swimming performance of Pagothenia borchgrevinki was unaffected by exposure to either -1 degrees C or 4 degrees C for 4 weeks. Maximum swimming velocity of both acclimation groups was thermally independent over the total temperature range of 1 degrees C to 10 degrees C. Therefore, the loss of any capacity to restructure the phenotype and an inability to thermally acclimate swimming performance appears to be associated with inhabiting a highly stable thermal environment.

Acclimatization↗

Impact of water temperature and stressor controllability on swim stress-induced changes in body temperature, serum corticosterone, and immobility in rats.

The present study compared the effects of three different water temperatures (20, 25, and 30 degrees C) and stressor controllability on several physiological and behavioral endpoints in an intermittent swim stress paradigm. The escape latency of rats in the 20 and 25 degrees C water was less than that observed for the 30 degrees C group. Both escape and yoked groups at 20 and 25 degrees C exhibited moderate to severe hypothermia following the swim stress session that returned to prestress levels 30-40 min post-stress. At 30 degrees C core body temperature (Tb) only decreased by 1 degree C for either swim group. Following swim, serum corticosterone (CORT) levels were significantly elevated in both escape and yoked groups in comparison to confined and home cage controls. The confined control group showed a significant elevation that was approximately halfway between the home cage control and the swim stress groups. At 30 degrees C, there was still a significant elevation of serum CORT in both swim groups in comparison to confined and home cage controls. Therefore, 30 degrees C appears to be the optimal water temperature to evaluate stress controllability effects in the current paradigm. In a final experiment, swim stressor controllability effects were examined in a 5 min forced swim test (FST) 24 h following the initial stress exposure. Rats exposed to yoked-inescapable swim stress at 30 degrees C exhibited more immobility than their escapable swim stress and confined counterparts, while the escape and confined controls did not differ. These results demonstrate that the behavioral deficits observed in the FST are attributable to the stress of inescapable swim and not swim stress per se.

Animals↗

The influence of temperature on muscle velocity and sustained performance in swimming carp.

The aim of this study was to evaluate how fish locomote at different muscle temperatures. Sarcomere length excursion and muscle shortening velocity, V, were determined from high-speed motion pictures of carp, Cyprinus carpio (11-14 cm), swimming steadily at various sustained speeds at 10, 15 and 20 degrees C. In the middle and posterior regions of the carp, sarcomeres of the lateral red muscle underwent cyclical excursions of 0.31 microns, centered around the resting length of 2.06 microns (i.e. from 1.91 to 2.22 microns). The amplitudes of the sarcomere length excursions were essentially independent of swimming speed and temperature. As tail-beat frequency increased linearly with swimming speed regardless of temperature, the sarcomeres underwent the same length changes in a shorter time. Thus, V increased in a linear and temperature-independent manner with swimming speed. Neither temperature nor swimming speed had an influence on tail-beat amplitude or tail height. Our findings indicate that muscle fibres are used only over a narrow, temperature-independent range of V/Vmax (0.17-0.36) where power and efficiency are maximal. Carp start to recruit their white muscles at swimming speeds where the red muscle V/Vmax becomes too high (and thus power output declines). When the V/Vmax of the active muscle falls too low during steady swimming, carp switch to 'burst-and-coast' swimming, apparently to keep V/Vmax high. Because Vmax (maximum velocity of shortening) of carp red muscle has a Q10 of 1.63, the transition speeds between swimming styles are lower at lower temperatures. Thus, carp recruit their white anaerobic muscle at a lower swimming speed at lower temperatures (verified by electromyography), resulting in a lower maximum sustainable swimming speed. The present findings also indicate that, to generate the same total force and power to swim at a given speed, carp at 10 degrees C must recruit about 50% greater fibre cross-sectional area than they do at 20 degrees C.

Animals↗

The influence of thermal acclimation on power production during swimming. II. Mechanics of scup red muscle under in vivo conditions.

We have previously shown that the power output of red muscle from warm-acclimated scup is greatly reduced when the fish swim at low temperatures. This reduction occurs primarily because, despite the slowing of muscle relaxation rate at cold temperatures, warm-acclimated scup swim with the same tail-beat frequency and the same stimulation durations, thereby not affording the slower-relaxing muscle any extra time to relax. We hypothesize that power output during swimming could be increased if the stimulus duration were reduced or if the relaxation rate of the red muscle were increased during cold acclimation. Scup were acclimated to 10 degrees C (cold-acclimated) and 20 degrees C (warm-acclimated) for at least 6 weeks. Cold acclimation dramatically increased the ability of scup red muscle to produce power at 10 degrees C. Power output measured from cold-acclimated muscle bundles driven through in vivo conditions measured from cold-acclimated scup swimming at 10 degrees C (i.e. work loops) was generally much greater than that from warm-acclimated muscle driven through its respective in vivo conditions at 10 degrees C. The magnitude of the increase depended both on the anatomical location of the muscle and on swimming speed. Integrated over the length of the fish, the red musculature from cold-acclimated fish generated 2.7, 8.9 and 5.8 times more power than the red musculature from warm-acclimated fish while swimming at 30 cm s(-)(1), 40 cm s(-)(1) and 50 cm s(-)(1), respectively. Our analysis suggests that the cold-acclimated fish should be able to swim in excess of 40 cm s(-)(1) with just their red muscle whereas the warm-acclimated fish must recruit their pink muscle well below this speed. Because the red muscle is more aerobic than the pink muscle, cold acclimation may increase the sustained swimming speed at which scup perform their long seasonal migrations at cool temperatures. We then explored the underlying mechanisms for the increase in muscle power output in cold-acclimated fish. Contrary to our expectations, cold-acclimated muscle did not have a faster relaxation rate; instead, it had an approximately 50 % faster activation rate. Our work-loop studies showed that this faster activation rate, alone, can increase the mechanical power production during cyclical contractions to a surprising extent. By driving cold-acclimated muscle through warm- and cold-acclimated in vivo conditions, we were able to partition the improvement in power production associated with increased activation rate and the approximately 20 % reduction in the duration of electromyographic activity found in the accompanying study. Depending on the position and swimming speed, approximately 60 % of the increase in power output was due to the change in the red muscle's contractile properties (i.e. faster activation); the remainder was due to the shorter stimulus duty cycle of cold-acclimated scup. Thus, by both shortening the in vivo stimulation duration and speeding up the rate of muscle activation as part of cold-acclimation, scup achieve a very large increase in the power output of their red muscle during swimming at low temperature. This increase in power output probably results in an increase in muscle efficiency and, hence, a reduction in the energetic cost of swimming. This increase in power output also reduces reliance on the less aerobic and less fatigue-resistant pink muscle. Both these abilities may increase the swimming speed at which prolonged aerobic muscle activity can occur and thus reduce the travel time for the long seasonal migrations in which scup engage.

Adaptation, Physiological↗

Patterns in early embryonic motility: effects of size and environmental temperature on vertical velocities of sinking and swimming echinoid blastulae.

Early embryonic swimming is widespread among marine invertebrates, but quantitative information about swimming behaviors is scarce. Swimming may affect encounters with predators, positioning in the water column, and nutrient absorption. Measured rates and patterns of swimming and sinking for blastulae of four eastern Pacific echinoid species show that sinking speeds equal or exceed swimming speeds. Swimming speed scaled negatively with embryo size, though sinking speed did not scale with size. Analysis of swimming paths of Strongylocentrotus franciscanus revealed a temperature dependency in swimming pattern that affected speed of upward movement. Sinking speeds were significantly greater at 10 degrees C than at 14 degrees C for blastulae of all four species examined. In Dendraster excentricus, killing the blastulae annulled this temperature effect, indicating an active density regulation by these embryos. Finally, measurements of particle velocities around sinking and swimming D. excentricus blastulae show that swimming creates a more localized disturbance than sinking. Embryonic swimming may therefore decrease rather than increase encounters with pelagic predators. Results from subsequent experiments in which embryos were reared in low-oxygen environments suggest that any oxygen-absorption advantages of swimming have little, if any, effect on the development of D. excentricus embryos.

Animals↗

The effect of repeated exposure to forced swimming on extracellular levels of 5-hydroxytryptamine in the rat.

The effects of repeated exposure to forced swimming was examined on extracellular concentrations of 5-hydroxytryptamine (5-HT), behavioral, and endocrine responses in rats. Animals were exposed to a 15-min swimming session on two consecutive days. On the first day, the swimming session increased extracellular 5-HT by 80 % over baseline in the striatum and reduced 5-HT to 40 % below baseline in the lateral septum. On the second day, however, the swimming session produced no effect on 5-HT in either brain region. Ratings of behavior showed that rats demonstrated climbing and swimming behaviors and developed immobility during the initial swimming session and that ratings of immobility increased and swimming decreased during the second swimming session. Immobility was positively correlated and swimming was negatively correlated with changes in extracellular 5-HT in the lateral septum but not in the striatum. Plasma corticosterone was equally elevated by 950 % after either 1 or 2 days of swimming exposure. These results show that there is rapid adaptation to the effects of repeated forced swimming on the regionally-specific, bi-directional response of extracellular 5-HT. In addition, changes in extracellular 5-HT in the lateral septum may be related to the behaviors produced during the forced swimming test that underlie its utility as an animal model of depression.

Animals↗