Locomotion in copepods: pattern of movements and energetics of Cyclops

The cost of swimming in copepods has generally been estimated through the application of fluid dynamics theory to data on velocity and acceleration obtained by means of movies. It has also been estimated through the changes in fat content of copepods after sustained swimming (i.e. vertical migration). However, the range of estimated costs of locomotion is exceedingly large (from 0.1% to 95% of total metabolism). This communication studies the pattern of swimming movements and the work done by Cyclops, using high speed cinematographic techniques. The contribution of swimming to the energy expenditure of the individual is estimated, and consideration of the possible role of rubber-like proteins in the cuticle of copepods is made.     

From Oxford to Hawaii ecophysiological barriers limit human progression in ten sport monuments

In order to understand the determinants and trends of human performance evolution, we analyzed ten outdoor events among the oldest and most popular in sports history. Best performances of the Oxford-Cambridge boat race (since 1836), the channel crossing in swimming (1875), the hour cycling record (1893), the Elfstedentocht speed skating race (1909), the cross country ski Vasaloppet (1922), the speed ski record (1930), the Streif down-hill in Kitzbühel (1947), the eastward and westward sailing transatlantic records (1960) and the triathlon Hawaii ironman (1978) all follow a similar evolutive pattern, best described through a piecewise exponential decaying model (r(2) = 0.95+/-0.07). The oldest events present highest progression curvature during their early phase. Performance asymptotic limits predicted from the model may be achieved in fourty years (2049+/-32 y). Prolonged progression may be anticipated in disciplines which further rely on technology such as sailing and cycling. Human progression in outdoor sports tends to asymptotic limits depending on physiological and environmental parameters and may temporarily benefit from further technological progresses.     

Effect of different warm-up procedures on subsequent swim and overall sprint distance triathlon performance

This study investigated the effect of 3 warm-up procedures on subsequent swimming and overall triathlon performance. Seven moderately trained, amateur triathletes completed 4 separate testing sessions comprising 1 swimming time trial (STT) and 3 sprint distance triathlons (SDT). Before each SDT, the athletes completed 1 of three 10-minute warm-up protocols including (a) a swim-only warm-up (SWU), (b) a run-swim warm-up (RSWU), and (c) a control trial of no warm-up (NWU). Each subsequent SDT included a 750-m swim, a 500-kJ (～20 km) ergometer cycle and a 5-km treadmill run, which the athletes performed at their perceived race intensity. Blood lactate, ratings of perceived exertion, core temperature, and heart rate were recorded over the course of each SDT, along with the measurement of swim speed, swim stroke rate, and swim stroke length. There were no significant differences in individual discipline split times or overall triathlon times between the NWU, SWU, and RSWU trials (p > 0.05). Furthermore, no difference existed between trials for any of the swimming variables measured (p > 0.05) nor did they significantly differ from the preliminary STT (p > 0.05). The findings of this study suggest that warming up before an SDT provides no additional benefit to subsequent swimming or overall triathlon performance.     

Swimming speed, respiration rate, and estimated cost of transport in adult killer whales

The physiology of free-ranging cetaceans is difficult to study and as a consequence, data on the energetics of these animals are limited. To better understand the energetic cost of swimming in killer whales, total cost of transport ( COT ) was estimated from swimming speeds and respiration rates from wild adult northern resident killer whales ( Orcinus orca ) and reported values of oxygen consumption in captive whales. Respiration rate (breaths per minute) was positively correlated with swimming speed (meters per second), while mass-specific COT (Joules per kilogram per meter) decreased with speed. Lack of data on very fast-swimming animals hindered assessment of the exact speed at which COT was minimal. However, minimum mass-specific COT for killer whales in the present study approached those predicted by a previously published allometric equation for marine mammals, and corresponded to "optimal" swimming speeds of 2.6–3 m/s. Interestingly, the observed average swimming speed (1.6 m/s) was lower than predicted optimal swimming speed. Finally, females with dependent calves had higher respiration rates than females without calves. These findings could be due to synchronous breathing with calves or could result from increased costs of lactation and swimming with a calf in echelon formation. Consequently, females with calves may have much greater COT at optimal swimming speeds than females without calves.     

Estimation of fish activity costs using underwater video cameras

Swimming speed and activity costs of dace ( Phoxinus eos × P. neogaeus ) were estimated in the field using underwater video cameras. Activity costs were estimated by converting swimming speeds and the number of movements into swimming costs. Average swimming speed ranged from 6.7 to 12.2 cm.s⁻¹ across 2 h periods and varied significantly among dates and time of day. The time spent swimming by dace ranged from 616 to 17 640 s 2 h⁻¹. Activity costs per 2h period ranged from 2.1 to 4O.2J 2h⁻¹ and were strongly correlated to the time spent swimming. Daily activity cost estimated using the cameras averaged 128.9J day⁻¹ and was equivalent to 1.7 times the standard metabolic rate. Activity cost predicted using a bioenergetic model in conjunction with independent estimates of consumption and growth rates averaged 138.8J day⁻¹. This study indicated that swimming characteristics and activity costs of dace varied significantly both within and among days. These analyses also indicated that equally valid activity costs for fish in the field can be estimated using video cameras and the difference between Consumption and growth rates.     

The use of computer-assisted motion analysis for quantitative studies of the behaviour of barnacle (Balanus amphitrite) larvae

The effects of larval density and age on pre-settlement swimming behaviour of Balanus amphitrite cyprid larvae were studied with the aid of computer-assisted motion analysis. Swimming behaviour was monitored in individual, in groups of 10-15 and in groups of 50-100 cyprids. There was a small, but significant effect of larval density on swimming speed and no effect on two other quantitative measures: rate of change of direction and net-to-gross displacement ratio. There was also small but significant variation in swimming speed between different batches of cyprids over the course of 2 years. Swimming behaviour of individual cyprid larvae was also monitored daily for 7 days, with the larvae maintained in the cold and dark between measurements to prevent settlement and metamorphosis. There were no significant behavioural differences observed over time indicating that larvae may be held in this manner experimentally without affecting these parameters.     

Simultaneous measurement of bacterial flagellar rotation rate and swimming speed.

Swimming speeds and flagellar rotation rates of individual free-swimming Vibrio alginolyticus cells were measured simultaneously by laser dark-field microscopy at 25, 30, and 35 degrees C. A roughly linear relation between swimming speed and flagellar rotation rate was observed. The ratio of swimming speed to flagellar rotation rate was 0.113 microns, which indicated that a cell progressed by 7% of pitch of flagellar helix during one flagellar rotation. At each temperature, however, swimming speed had a tendency to saturate at high flagellar rotation rate. That is, the cell with a faster-rotating flagellum did not always swim faster. To analyze the bacterial motion, we proposed a model in which the torque characteristics of the flagellar motor were considered. The model could be analytically solved, and it qualitatively explained the experimental results. The discrepancy between the experimental and the calculated ratios of swimming speed to flagellar rotation rate was about 20%. The apparent saturation in swimming speed was considered to be caused by shorter flagella that rotated faster but produced less propelling force.     

Bioenergetic model of planktivorous fish feeding, growth and metabolism: theoretical optimum swimming speed of fish larvae

The feeding activity of an individual fish larva is described by an equation which includes parameters for the area successfully searched, probability of food capture multiplied by the cross-sectional perceptive visual field, larval swimming speed and the time required to consume a unit of food energy. The proportion of ingested food energy used for metabolism increases exponentially with increasing swimming speed. The model predicts that food consumption rate increases asymptotically whereas metabolic rate increases exponentially. This results in a predicted growth rate curve that reaches a maximum at a certain swimming speed and decreases at both higher and lower speeds. The model can be used to predict the influence of type of prey, prey density, water temperature etc. on larval growth. An expression describing how many hours per day fish larvae must forage in order to grow at a certain daily body weight gain allows the limits of environmental conditions for positive, zero and negative growth rate to be set. Results of simulations demonstrated that the optimum swimming speed for maximum growth of coregonid larvae increased with an increase in food density, decrease in water temperature or decrease of prey vulnerability. At optimum ‘theoretical’ swimming speed an increase in water temperature from 5 to 17° C required the food density to be increased from 20 to 80 copepods l^?1 in order to maintain a daily growth increment of 2%. The minimum Artemia density required for maintenance metabolism increased from 10 to 30 items 1¹ over the same temperature increase from 5 to 17° C, and food densities required for 8% growth rates were 26 and 56 Artemia nauplii l^?1 at 5 and 17° C, respectively. Contrary to previous findings, results of the present study suggest that metabolic rates of actively feeding fish larvae may be from 5 to 50 times the standard metabolic rate: earlier studies suggested that a factor of 2–3 may be generally applicable.     

Effect of position-fixing interval on estimated swimming speed and movement pattern of fish tracked with a stationary positioning system

Ultrasonic telemetry using stationary positioning systems allows several fish to be tracked simultaneously, but systems that are incapable of sampling multiple frequencies simultaneously can record data from only one transmitter (individual) at a time. Tracking several individuals simultaneously thus results in longer intervals between successive position fixes for each fish. This deficiency leads to loss of detail in the tracking data collected, and may be expected to cause loss of accuracy in estimates of the swimming speeds and movement patterns of the fish tracked. Even systems that track fish on multiple frequencies are not capable of continuous tracking due to technical issues. We determined the swimming speed, area occupied, activity rhythm and movement pattern of cod (Gadus morhua) using a stationary single-channel positioning system, and analysed how estimates of these behavioural parameters were affected by the interval between successive position fixes. Single fish were tracked at a time, and position fixes were eliminated at regular intervals in the original data to generate new data sets, as if they had been collected in the course of tracking several fish (2–16). In comparison with the complete set, these data sets gave 30–70% decreases in estimates of swimming speed depending on the number of fish supposedly being tracked. These results were similar for two individuals of different size and activity level, indicating that they can be employed as correction factors to partly compensate for underestimates of swimming speed when several fish are tracked simultaneously. Tracking `several' fish only slightly affected the estimates of area occupied (1–15%). The diurnal activity rhythm was also similar between the data sets, whereas details in search pattern were not seen when several fish were tracked simultaneously.     

Rotation and switching of the flagellar motor assembly in Halobacterium halobium.

Halobacterium halobium swims with a polarly inserted motor-driven flagellar bundle. The swimming direction of the cell can be reserved by switching the rotational sense of the bundle. The switch is under the control of photoreceptor and chemoreceptor proteins that act through a branched signal chain. The swimming behavior of the cells and the switching process of the flagellar bundle were investigated with a computer-assisted motion analysis system. The cells were shown to swim faster by clockwise than by counterclockwise rotation of the flagellar bundle. From the small magnitude of speed fluctuations, it is concluded that the majority, if not all, of the individual flagellar motors of a cell rotate in the same direction at any given time. After stimulation with light (blue light pulse or orange light step-down), the cells continued swimming with almost constant speed but then slowed before they reversed direction. The cells passed through a pausing state during the change of the rotational sense of the flagellar bundle and then exhibited a transient acceleration. Both the average length of the pausing period and the transient acceleration were independent of the stimulus size and thus represent intrinsic properties of the flagellar motor assembly. The average length of the pausing period of individual cells, however, was not constant. The time course of the probability for spontaneous motor switching was calculated from frequency distribution and shown to be independent of the rotational sense. The time course further characterizes spontaneous switching as a stochastic rather than an oscillator-triggered event.     

Maximum sustainable swimming speeds of late-stage larvae of nine species of reef fishes

We examined the maximum sustainable swimming speed of late-stage larvae of nine species of tropical reef fishes from around Lizard Island, Great Barrier Reef, Australia. Larvae were captured in light traps and were swum in flumes at different experimental swimming speeds (of 5 cm s⁻¹ intervals) continuously for 24 h. Logistic regression was used to determine the speed at which 90% of larvae were able to maintain swimming, and this was used to indicate the maximum sustainable swimming speed for each species. Maximum sustainable swimming speeds varied among the species examined, with the lethrinid maintaining the fastest sustainable swimming speed (24 cm s⁻¹), followed by the Pomacentridae (10-20 cm s⁻¹) and the Apogonidae (8-12 cm s⁻¹). U-crit (maximum speed) explained 64% of the variation in sustainable speed among species, whereas total length only explained 33% of the variation in sustained swimming. A regression fitted across species suggests that 50% U-crit is a good approximation of the speed able to be maintained by these larvae for 24 h. A model based on a cubic relationship between sustained swimming time and speed was found to be more successful than either length or U-crit as a method of estimating sustainable swimming speed for most of the species examined. Overall, we found that swimming speed is an important factor when considering the potential for active swimming behaviour to influence dispersal patterns, recruitment success and levels of self-recruitment in reef fish larvae and needs to be carefully considered in models of larval dispersal.     

Motility of bovine spermatozoa studied by laser light scattering

Laser light scattering has been employed to determine the swimming speed distribution and the fraction of motile cells in samples of bovine spermatozoa. As predicted from theory, average trajectory velocities determined by laser light scattering were approximately four times the average translational speed estimated using light microscopy. The proportion of motile spermatozoa decreased with time at the same rate when samples were prepared in either HEPES or phosphate buffers. However, whereas the mean swimming velocity declined slowly in HEPES buffer, it dropped rapidly when phosphate buffer was used. Dilution (in the range 40–0.4×10⁶ spermatozoa·ml^-1) in either of these two buffers reduced the fraction of motile spermatozoa in the sample, but the mean swimming velocity of the remaining active spermatozoa was unchanged. Lowering the temperature from 37° C to 15° C reduced the mean swimming speed by a factor of 2–3 and the fraction of motile cells by a factor of 4–5.     

Seasonal analysis of Daphnia pulicaria swimming behavior

Our study documents individual swimming behavior of Daphnia pulicaria over a yearly cycle in a temperate lake. We collected D. pulicaria, a common freshwater zooplankton, from Lake Mendota on 10 dates between July 1994 and June 1995 from two depths, 2 m and 10 m. The Daphnia were rushed to the laboratory and video-taped as they swam in lake water under lake-ambient temperature and light conditions. Five-second swimming tracks of individual Daphnia were filmed and digitized using a motion analysis system. We measured average turning angle, swimming speed and sinking rate for each track. D. pulicaria swimming behavior varied over the annual cycle. We found significant differences in turning angle between depths and among months. Sinking rate and swimming speed were significantly different among months but not depths. Sinking rate and swimming speed were not significantly correlated with water temperature. Our results were contrary to Stokes' Law predictions, in that D. pulicaria had the slowest sinking speed in June, not in the winter when water temperatures were lowest and viscosity was highest. Body length was significantly correlated with all three swimming variables. We also studied swimming behavior in clonal populations of D. pulicaria in different concentrations of the alga, Chlamydomonas reinhardtii. D. pulicaria did not change swimming speed, turning angle or sinking rate over a range of food concentrations. Finally, swimming behavior in a D. pulicaria clone, tested at two temperatures in the laboratory, confirmed the results from our seasonal study; Daphnia did not sink as predicted by changes in viscosity.     

不同程度的尾损伤对镇海林蛙(Rana zhenhaiensis)蝌蚪游泳速度的影响

无尾两栖类蝌蚪的尾巴通过产生强大的游泳速度在反捕食中起到了重要的作用。以镇海林蛙(Rana zhenhaiensis)蝌蚪为实验动物来评估断尾的运动代价。以74尾具有完整尾蝌蚪作为实验组,通过截去不同尾长片段,人为分成轻微尾损伤组(30%)和严重尾损伤组(30%)并测定两组蝌蚪在断尾前后的游泳速度。以16尾完整尾蝌蚪作为对照组在实验组断尾前后同时进行游泳速度的测定。实验结果显示断尾影响蝌蚪的游泳速度,但仅在尾损伤程度达到尾长的30%以上时才产生不利的影响。这表明轻微尾损伤并不对镇海林蛙蝌蚪的游泳速度产生严重影响。在断尾前后实验组蝌蚪的游泳速度均与尾长呈正相关。在相同尾长状态下,尾损伤蝌蚪的相对游泳速度明显快于完整尾蝌蚪。因此,尾损伤的镇海林蛙蝌蚪有可能通过改变尾和身体的摆动频次等方式在断尾后对游泳速度进行了一定的补偿。尾损伤在野外频繁发生于蝌蚪的尾远端,据此推测镇海林蛙蝌蚪在自然条件下的尾损伤并不会产生严重运动代价。     

A study of the swimming performance of the Crucian carp Carassius carassius (L.) in relation to the effects of exercise and recovery on biochemical changes in the myotomal muscles and liver

A study has been made of the maximum sustained swimming speed of Crucian carp Carassius carassius (L.) using a fixed velocity technique. The data obtained from swimming tests on 214 carp have been analysed using the method of probit analysis. The 50% fatigue level for 13–16 cm fish acclimated to 9.5±0.6°C has been estimated to be 3.35 lengths/sec. Biochemical measurements have been made on the red and white myotomal muscles and liver of fish subjected to both varying intensities of sustained swimming and short periods of vigorous swimming. Free creatine was found to increase only during high speed swimming in the white muscle. Elevated lactate concentrations occurred at both low and high sustained swimming speeds in the red superficial muscle but not during short periods of strenuous exercise. Glycogen depletion from the red musculature also only took place at the sustained swimming speeds investigated. The reverse situation was operative in the white muscle, significant glycogen depletion occurring only at the highest swimming speed studied. Lactate levels were only significantly different from non-exercised fish in the fish swimming at the higher velocities. The effects of periods of recovery following 200 min of sustained swimming were also investigated. White muscle lactate was at a higher level than non-exercise fish 5 h post-exercise, while both red muscle glycogen and lactate rapidly returned to pre-exercise concentrations. Biochemical measurements on the myotomal muscle types have been discussed in relation to the swimming performance of the fish and the division of labour between red and white fibres.     

Behaviour of rainbow trout Oncorhynchus mykiss presented with a choice of normoxia and stepwise progressive hypoxia

The objective of this study was to identify behavioural adjustments leading to avoidance of hypoxia. Using the oxygen-sensitive species rainbow trout Oncorhynchus mykiss as a model, individual fish were recorded while moving freely between two sides of a test arena: one with normoxia and one with stepwise progressive hypoxia [80-30% dissolved oxygen (DO) air saturation]. The results demonstrated a gradual decrease in the total time spent in hypoxia starting at 80% DO air saturation. At this DO level, the avoidance of hypoxia could not be attributed to changes in spontaneous swimming speed, neither in normoxia nor in hypoxia. Reducing the DO level to 60% air saturation resulted in decreased spontaneous swimming speed in normoxia, yet the number of trips to the hypoxic side of the test arena remained unchanged. Moreover, data revealed increased average residence time per trip in normoxia at DO levels ≤60% air saturation and decreased average residence time per trip in hypoxia at DO levels ≤50% air saturation. Finally, the spontaneous swimming speed in hypoxia increased at DO levels ≤40% air saturation and the number of trips to hypoxia decreased at the 30% DO air saturation level. Thus, avoidance of the deepest hypoxia was connected with a reduced number of trips to hypoxia as well as decreased and increased spontaneous swimming speed in normoxia and hypoxia, respectively. Collectively, the data support the conclusions that the mechanistic basis for avoidance of hypoxia may (1) not involve changes in swimming speed during mild hypoxia and (2) depend on the severity of hypoxia.     

Living in the fast lane: effects of cost of locomotion on foraging behaviour in juvenile Atlantic salmon

Stream-dwelling, juvenile Atlantic salmon, Salmo salar L., feed mainly on drifting invertebrates, usually by swimming upstream from a stationary position to intercept individual prey items. Laboratory experiments tested the prediction that individual salmon should reduce the distance over which they would travel (attack distance) to intercept drifting food items as the energy cost of swimming increases with increasing current velocity. Attack distance varied inversely with current velocity as expected. The fish's average speed of upstream movement relative to the substrate remained constant and the duration of individual attacks therefore declined as current velocity increased. Calculated reaction distances and a second ecperiment using tethered prey drifting at speeds independent of current velocity confirmed that these relationships were due to fish actually delaying attacks on perceived prey for longer periods as current velocity increased. Using estimated metabolic rates for burst swimming, it appears that energy expenditure per attack varies little with current velocity. Therefore, by reducing their reaction and attack distances in response to increasing current velocity, the fish reduced their energy cost of travel per attack.     

Activity patterns of juvenile Atlantic cod (Gadus morhua) in Buckley Cove,Newfoundland

We monitored swimming speed of 2–3 year-old juvenile Atlantic cod (Gadus morhua) from August to December 1999, using a 2-D location finding acoustic telemetry system in a coastal area of Newfoundland, Canada. We concurrently monitored the locations of 22–41 individuals by triangulation using a fixed hydrophone array. We estimated average swimming speeds at intervals of 60–120 s and compared them over a 1 to 17 °C thermal range, three diel periods, and five substrates (sand, gravel, sand-sparse boulder, boulder, and kelp). However, cod did not exhibit a change in swimming speed over the temperature range studied. Increased activity and foraging rates (expressed as swimming speeds) were expected to increase at elevated temperatures due to increased metabolic demands. Activity did vary significantly with diel cycle and substrate. Swimming speeds were significantly lower at night during September and October. Results for August and November were inconclusive, while swimming speed was significantly lower during the day in December. We observed significantly reduced average swimming speeds in structurally complex substrates (e.g. rock, cobble and kelp) in September and October. Our results suggest that activity of juvenile cod in the wild does not vary with temperature as predicted from studies in the laboratory. Instead, activity varied with diel cycles and structural complexity, variables that influence an individual's ability to forage and seek refuge, potentially altering individual fitness.     

Tilting behaviour of the Atlantic mackerel, Scomber scombrus, at low swimming speeds

Negatively-buoyant Atlantic mackerel, Scomber scombrus L., (fork length 30–39 cm) tilt their bodies with the head up while swimming at speeds below 0.8 body length per second (B.L. s⁻¹). This behaviour is quantitatively described by the body attack angle and swimming speed measured from film records. The maximum recorded body attack angle was 27° in a 32 cm-long fish swimming at 0.45 B.L. s⁻¹ while its nose followed a course close to the horizontal. In general, larger body attack angles were shown at lower swimming speeds and were associated with denser bodies at each speed. We consider that this behaviour pattern allows the fish to maintain a chosen swimming depth while its body creates lift by acting as a hydrofoil. Lift from the fins is insufficient at low swimming speeds.     

Upstroke Thrust, Drag Effects, and Stroke-glide Cycles in Wing-propelled Swimming by Birds

A number of bird species swim underwater by wing propulsion.Both among and within species, thrust generated during the recoveryphase (upstroke) varies from almost none to more than duringthe power phase (downstroke). More uneven thrust and unsteadyspeed may increase swimming costs because of greater inertialwork to accelerate the body fuselage (head and trunk), especiallywhen buoyant resistance is high during descent. I investigatedthese effects by varying relative fuselage speed during upstrokevs. downstroke in a model for wing-propelled murres which descendat relatively constant mean speed. As buoyant resistance declinedwith depth, the model varied stroke frequency and glide durationto maintain constant mean descent speed, stroke duration, andwork per stroke. When mean fuselage speed during the upstrokewas only 18% of that during the downstroke, stroke frequencywas constant with no gliding, so that power output was unchangedthroughout descent. When mean upstroke speed of the fuselagewas raised to 40% and 73% of mean downstroke speed, stroke frequencydeclined and gliding increased, so that power output decreasedrapidly with increasing depth. Greater inertial work with moreunequal fuselage speeds was a minor contributor to differencesin swimming costs. Instead, lower speeds during upstrokes requiredhigher speeds during downstrokes to maintain the same mean speed,resulting in nonlinear increases in drag at greater fuselagespeeds during the power phase. When fuselage speed was relativelyhigher during upstrokes, lower net drag at the same mean speedincreased the ability to glide between strokes, thereby decreasingthe cost of swimming.