The swimming endurance of threespine sticklebacks, Gasterosteus aculeatus L., from the Afon Rheidol, Wales

The endurance of threespine sticklebacks, Gasterosteus aculeatus , swimming with pectoral fin locomotion at 20° C in a laboratory flume was measured. Each trial lasted a maximum of 480 min. At a speed of 4 body lengths per sec (L s⁻¹) all fish were still swimming at the end of the trial, but endurance decreased at higher speeds. At speeds of 5 or 6 L s⁻¹ (20–30 cm s⁻¹) a few fish still maintained labriform locomotion for the 480 min. However, at a speed of 7 L s⁻¹ all fish furled their pectoral fins and used body and caudal fin propulsion but fatigued rapidly. During sustained swimming, fish could cover distances of 6 km or more. No significant differences between males and females were found.     

Is tilting behaviour at low swimming speeds unique to negatively buoyant fish? Observations on steelhead trout,Oncorhynchus mykiss,and bluegill,Lepomis macrochirus

When swimming at low speeds, steelhead trout and bluegill sunfish tilted the body at an angle to the mean swimming direction. Trout swam using continuous body/caudal fin undulation, with a positive (head-up) tilt angle ( 0 , degrees) that decreased with swimming speed ( u , cm s⁻¹) according to: 0 =(164±96).u^{(−1.14±0.41)} (regression coefficients; mean±2 s.e. ). Bluegill swimming gaits were more diverse and negative (head down) tilt angles were usual. Tilt angle was −3·0 ± 0.9° in pectoral fin swimming at speeds of approximately 0.2–1.7 body length s⁻¹ (Ls⁻¹; 3–24 cm s⁻¹), −4.5 ±2.6° during pectoral fin plus body/caudal fin swimming at 1·2–1·7 L s⁻¹ (17–24cm s⁻¹), and −5.0± 1.0° during continuous body/caudal fin swimming at 1.6 and 2.5 L s⁻¹ (22 and 35cm s⁻¹). At higher speeds, bluegill used burst-and-coast swimming for which the tilt angle was 0.1±0.6°. These observations suggest that tilting is a general phenomenon of low speed swimming at which stabilizers lose their effectiveness. Tilting is interpreted as an active compensatory mechanism associated with increased drag and concomitant increased propulsor velocities to provide better stabilizing forces. Increased drag associated with trimming also explains the well-known observation that the relationship between tail-beat frequency and swimming speed does not pass through the origin. Energy dissipated because of the drag increases at low swimming speeds is presumably smaller than that which would occur with unstable swimming.     

The muscle twitch and the maximum swimming speed of giant bluefin tuna, Thunnus thynnus L.

Sustained swimming of bluefin tuna was analysed from video recordings made of a captive patrolling fish school [lengths (L) 1.7–3.3 m, body mass (M) 54–433 kg]. Speeds ranged from 0.6 to 1.2 L s⁻¹ (86–260 km day⁻¹) while stride length during steady speed swimming varied between 0.54 and 0.93 L. Maximum swimming speed was estimated by measuring twitch contraction of the anaerobic swimming muscle in pithed fish 5 min after death. Muscle contraction time increased from the shortest just behind the head (30–50 ms at 20% L) to the longest at the tail peduncle (80–90 ms at 80% L) (all at 28°C). A fish (L = 2.26 m) with a muscle contraction time of 50 ms at 25% L can have a maximum tail beat frequency of 10 Hz and maximum swimming speed of 15m s⁻¹ (54km h⁻¹) with a stride length of 0.65L. With a stride length of 1 L a speed of 22.6 m s⁻¹ (81.4 km h⁻¹) is possible. Power used at maximum speed was estimated for this fish at between 10 and 40 kW, with corresponding values for the drag coefficient at a Reynolds number of 4.43 × 10⁷ of 0.0007 and 0.0027.     

Burst swimming speeds of mackerel, Scomber scombrus L.

Burst swimming speeds were measured in mackerel 0.275–0.380 m long by filming newly caught fish, first released into a large shore-sited tank, using a high-speed cine camera and real time TV camera. The highest speed was 5.50 m s⁻¹ or 18 body length per second ( b.l . s⁻¹) in a 0.305 m long mackerel at 12° C. The recorded maximum tail beat frequency of 18 Hz agrees well with 19 Hz predicted from the measured contraction time of 0.026 s for the anterior lateral swimming muscle. The stride length was close to 1 B.L.; the power, calculated from the drag, was 4.53 W, and, calculated from the muscle used, was 5.07 W; all suggesting that the mackerel is swimming close to its physiological limit.     

Swimming performance of European eel (Anguilla anguilla (L.)) elvers

To determine the relation between swimming endurance time and burst swimming speed, elvers of the European eel, Anguilla anguilla (L.), were made to swim at speeds from 3.6 to 7.2 L (body lengths) s⁻¹ in both fresh and sea water. Swimming endurance time of elvers averaging 7.2 cm total length decreased logarithmically with increased swimming speed from 3.0 min at 3.5 L s⁻¹ to 0.7 min at 5.0 L s⁻¹, and again logarithmically but with a lesser slope to 0.27 min at 7.5 L s⁻¹. No differences were found between fresh and sea water elvers. In still water, elvers could swim at high speeds for about 10–45m before exhaustion, depending upon speed. Elvers would be able to make virtually no progress against water currents >50 cm s⁻¹. Drift in coastal water currents and selective tidal transport probably involve swimming speeds below those tested in this study. Migration into freshwater streams undoubtedly involves avoidance of free stream speeds and a combination of burst and sustained swimming.     

Endurance swimming of diploid and triploid Atlantic salmon

When groups of diploid (mean ±  s . e . fork length, L _F) 33·0 ± 1·4 cm and triploid (35·3 ± 0·5 cm) Atlantic salmon Salmo salar were forced to swim at controlled speeds in a carefully monitored 10 m diameter 'annular' tank no significant difference was found between the maximum sustained swimming speeds ( U _ms, maintainable for 200 min) where the fish swam at the limit of their aerobic capability. Diploids achieved 2·99 body lengths per second (bl s⁻¹)(0·96 m s⁻¹) and triploids sustained 2·91 bl s⁻¹(1·02 m s⁻¹). The selection of fish for the trials was based on their ability to swim with a moving pattern projected from a gantry rotating at the radius of the tank and the selection procedure did not prove to be significant by ploidy. A significant difference was found between the anaerobic capabilities of the fish measured as endurance times at their prolonged swimming speeds. During the course of the experimentation the voluntary swimming speed selected by the fish increased and the schooling behaviour improved. The effect of the curvature of the tank on the fish speeds was calculated (removing the curved effect of the tank increased the speed in either ploidy by 5·5%). Implications of the endurance times and speeds are discussed with reference to the aquaculture of triploid Atlantic salmon.     

Hydroacoustic measurement of swimming speed of North Sea saithe in the field

Saithe Pollachius virens , tracked diurnally with a split-beam echosounder, showed no relationship between size and swimming speed. The average and the median swimming speeds were 1·05 m s⁻¹(±0·09 m s⁻¹) and 0·93 m s⁻¹, respectively. However, ping-to-ping speeds up to 3·34 m s⁻¹ were measured for 25–29 cm fish, whose swimming speeds were significantly higher at night (1·08 m s⁻¹) than during the day (0·72 m s⁻¹). The high average swimming speed could be related to the foraging or streaming part of the population and not to potential weakness of the methodology. However, the uncertainty of target location increased with depth and resulted in calculated average swimming speeds of 0·15 m s⁻¹ even for a stationary target. With increasing swimming speed the average error decreased to 0 m s⁻¹ for speeds >0·34 m s⁻¹. Species identity was verified by trawling in a pelagic layer and on the bottom.     

Constraints of body size and swimming velocity on the ability of juvenile rainbow trout to endure periods without food

The hypothesis that body size and swimming velocity affect proximate body composition, wet mass and size‐selective mortality of fasted fish was evaluated using small (107 mm mean total length, L _T) and medium (168 mm mean L _T) juvenile rainbow trout Oncorhynchus mykiss that were sedentary or swimming ( c . 1 or 2 body length s⁻¹) and fasted for 147 days. The initial amount of energy reserves in the bodies of fish varied with L _T. Initially having less lipid mass and relatively higher mass‐specific metabolic rates caused small rainbow trout that were sedentary to die of starvation sooner and more frequently than medium‐length fish that were sedentary. Swimming at 2 body length s⁻¹ slightly increased the rate of lipid catabolism relative to 1 body length s⁻¹, but did not increase the occurrence of mortality among medium fish. Death from starvation occurred when fish had <3·2% lipid remaining in their bodies. Juvenile rainbow trout endured long periods without food, but their ability to resist death from starvation was limited by their length and initial lipid reserves.     

Swimming performance and metabolism of 0+ year Thymallus arcticus

The prolonged swimming speed and metabolic rate of 0+ year Arctic grayling Thymallus articus were examined with respect to current velocity, water temperature and fish size, and compared to conditions fish occupy in the river. Oxygen consumption (mg O₂ h⁻¹) increased with fish mass and temperature (6–23° C), with a steep increase in metabolic rate between 12 and 16° C. Absolute prolonged swimming speed (cm s⁻¹) increased rapidly with fish size (total length, L _T, and mass), however, fish in the natural stream habitat occupied current velocities between 15 and 25 cm s⁻¹ or 4  L _T s⁻¹, approximately half their potential prolonged swimming speed (10  L _T s⁻¹).     

A hydro-mechanical approach to the scaling of swimming performance in the sand flathead Platycephalus bassensis Cuvier: effects of changes in morphological features based on fish size

The swimming performance of Platycephalus bassensis at steady speed was assessed with an emphasis on hydrodynamics. The minimum swimming speed to maintain hydrostatic equilibrium for P. bassensis of 0·271 m total length ( L _T) was calculated to be 1·06 L _T s⁻¹. At this speed, the required lift to support the mass of the fish was equivalent to 6·6% of the fish mass; 82·7% of which was created by the body as a hydrofoil, and the rest of which was created by the pelvic fins as hydrofoils. The minimum swimming speed decreased with the L _T of the fish and ranged from 1·15 L _T s⁻¹ for a fish of 0·209 m to 0·89 L _T s⁻¹ for a fish of 0·407 m. The forward movement per tail-beat cycle ( i.e. stride length) was described with an equation including quantities of morphological and hydro-mechanical relevance. This equation explained that stride length was increased by the effect of turbulence characterized by the Reynolds number and demonstrated the morphological and hydro-mechanical functional design of the fish for maximizing thrust and minimizing drag. The larger span of the caudal fin and caudal tail-beat amplitude was associated with larger stride length, whereas greater frictional drag was associated with smaller stride length.     

Behaviour and performance of juvenile shortnose sturgeon Acipenser brevirostrum at different water velocities

Critical swimming speeds (mean ± s . e .) for juvenile shortnose sturgeon Acipenser brevirostrum were 34·4 cm s⁻¹± 1·7 (2·18 ± 0·09 body lengths, BL s⁻¹). Swimming challenges at 10, 20 and 30 cm s⁻¹ revealed that juvenile A. brevirostrum are relatively poor swimmers, and that the fish did not significantly modify their swimming behaviour, although they spent more time substratum skimming ( i.e. contact with flume floor) at 30 cm s⁻¹ relative to 10 cm s⁻¹. When present, these behavioural responses are probably related to morphological features, such as flattened rostrum, large pectoral fins, flattened body shape and heterocercal tail, and may be important to reduce the costs of swimming.     

Circadian rhythms in juvenile american shad, Alosa sapidissima

Swimming activity (in cm s⁻¹) of a school (55 individuals) of young-of-the-year ( total length=110 mm) American shad, Alosa sapidissima , was determined under a variety of photoperiod conditions. These included a normal (ambient), a shifted, and constant-light day. Swimming activity was measured over 4-day periods. During normal days swimming speeds followed periods of about 24 h, with fast speeds (up to 45 cm s⁻¹) and schooling occurring during the photoperiod. Under dark conditions speeds were slower (8 cm s⁻¹) with most fish swimming as individuals. During a shifted day swimming speeds and schooling corresponded to the imposed day. Under constant light (equivalent to bright moonlight) no schooling was evident, and a constant, but slow, swimming speed was observed in each 24-h period. These shad demonstrated an exogenous rhythm with respect to the imposed day length. It is hypothesized that an endogenous circadian rhythm would only be of use to a fish required to hunt or chase its prey. Shad, being plankton feeders, do not chase prey and therefore can exhibit an exogenous circadian rhythm with no detrimental feeding results.     

Effects of the brain parasite Myxobolus arcticus on sockeye salmon

Parasitism with Myxobolus arcticus did not affect smolt size of sockeye salmon or their osmocompetence, but had a deleterious effect ( P <0.001) on the swimming speed of naturally infected smolts. Parasitized fish had a mean swimming speed of 2.89 fork length s⁻¹ (L_F s⁻¹) compared with 4.37 L _F s⁻¹ for unparasitized fish. The parasite probably impairs swimming ability by affecting the central nervous system, but this effect does not appear severe enough to limit the parasite's usefulness in stock separation.     

Swimming performance of juvenile sprat, Sprattus sprattus L., and herring, Clupea harengus L., at different salinities

Sustained swimming performance of juvenile sprat, S. sprattus (29–48 mm s.l.), and herring, C. harengus (46–58 mm) was measured in a laboratory flume over a range of salinities from 18 to 33%0 at water temperatures of 16–19°C. Critical swimming speeds (CSS) of both species, relative to body length, were similar, averaging 10–12 body lengths per second (bl s⁻¹). There was no apparent relationship with salinity.
These swimming speeds are higher than values generally quoted in the literature for sustained swimming of sprat and herring (2–7 bl s⁻¹) and it is concluded that the better performance found in this study was a function of improved fish handling techniques, and of the size of fish used since most other studies have dealt with larger, commercial sized fish.     

Gait transition and oxygen consumption in swimming striped surfperch Embiotoca lateralis Agassiz

A flow-through respirometer and swim tunnel was used to estimate the gait transition speed ( U _p-c) of striped surfperch Embiotoca lateralis , a labriform swimmer, and to investigate metabolic costs associated with gait transition. The U _p-c was defined as the lowest speed at which fish decrease the use of pectoral fins significantly. While the tail was first recruited for manoeuvring at relatively low swimming speeds, the use of the tail at these low speeds [as low as 0·75 body (fork) lengths s⁻¹, L _F s⁻¹) was rare (<10% of the total time). Tail movements at these low speeds appeared to be associated with occasional slow manoeuvres rather than providing power. As speed was increased beyond U _p-c, pectoral fin (PF) frequencies kept increasing when the tail was not used, while they did not when PF locomotion was aided by the tail. At these high speeds, the tail was employed for 40–50% of the time, either in addition to pectoral fins or during burst-and-coast mode. Oxygen consumption increased exponentially with swimming speeds up to gait transition, and then levelled off. Similarly, cost of transport ( C _T) decreased with increasing speed, and then levelled off near U _p-c. When speeds ≥ U _p-c are considered, C _T is higher than the theoretical curve extrapolated for PF swimming, suggesting that PF swimming appears to be higher energetically less costly than undulatory swimming using the tail.     

Importance of electrode positioning in biotelemetry studies estimating muscle activity in fish

Red and white axial muscle activity of adult Atlantic salmon Salmo salar was examined using conventional electromyography (EMG x ) and activity radio-transmitters (EMG i ) at 0·5 and 0.7 body lengths (L) along the body of the fish. Critical swimming trials were conducted and maximum sustainable speeds (U_crit) were unaffected by the presence of electrodes, being 1·51 ± 21 m s⁻¹ (3.33 ± 0.34 L s⁻¹) ( n =44). Regardless of longitudinal position of the electrodes within the musculature, both EMG x s and EMG i s indicated increasing red muscle activity with increasing swimming speed, whereas white muscle fibres were recruited only at speeds > 86±5% U_crit. Telemetered EMG i signals indicated that muscle activity varied significantly for electrodes implanted at different longitudinal positions along the fish ( P < 0·001). These results suggest that electrode placement is an important influence affecting the signals obtained from radio transmitters that estimate activity and location should be standardized within biotelemetry studies to allow accurate and consistent comparisons of activity between individuals and species. Optimal location for electrode placement was determined to be in the red muscle, towards the tail of the fish (0·7 L ).     

Sustained swimming speeds and myotomal muscle function in the trout, Salmo gairdneri

Rainbow trout were trained for 3–4 weeks in a flume at swimming speeds of 1, 2 and 3 l s⁻¹. For each experiment growth rates were estimated and by measuring the hypertrophy of red and mosaic skeletal muscle fibres their function was described at particular swimming speeds and compared with earlier experiments on coalfish using the same technique.
Maximum growth, compared with controls in still water, occurred at swimming speeds of 1 l s⁻¹. At this speed the trout mosaic muscle fibres hypertrophied by 40% but the red muscle fibres showed only a 25% hypertrophy. It is suggested that natural swimming speeds are close to 1Ls^−l and the trout mosaic fibres are better adapted for use at this speed in comparison with coalfish white muscle fibres.     

The critical swimming speed of the sand smelt (Atherina presbyter Cuvier) in relation to capture at a power station cooling water intake

Critical swimming speeds (CSS) of sand smelt, Atherina presbyter , were measured in a laboratory flume. Individuals of all age classes (0+ to III +) found in the vicinity of Fawley power station, Hampshire, were tested at temperatures covering the seasonal range. The median CSS was 2.7 body lengths per second (bl s⁻¹) at 5.8 °C, rising to 5.7 bl s⁻¹ at 18.5°C.
A two-variable (water temperature and body length) regression model was fitted to the data, and this was used to assess escape potential of fish coming into contact with the power station cooling water intake currents and the extent of possible bias in fish length data collected from this source. It is concluded that the sand smelt remains vulnerable to entrainment at the power station over its whole length range and over the full range of seasonal temperatures and that size-dependent swimming performance will not lead to significant bias in sample length distributions.     

The effect of external dummy transmitters on oxygen consumption and performance of swimming Atlantic cod

Decreased critical swimming speed and increased oxygen consumption (     ) was found for externally tagged Atlantic cod Gadus morhua swimming at a high speed of 0·9 body length (total length, L _T) s⁻¹. No difference was found in the standard metabolic rate, indicating that the higher     for tagged cod was due to drag force rather than increased costs to keep buoyancy.     

Influence of spawning on swimming performance and muscle contractile properties in the short-horn sculpin

The total distance travelled during the first two kinematic stages of the escape response of short-horn sculpin was significantly greater in post spawning fish (0·41 L) than in gravid fish (0·23 L). The maximum velocity of the snout during the C-bend was significantly higher (5·6 L s⁻¹) in postspawning fish than in gravid fish (3·8 L s⁻¹). To investigate some of the mechanisms underlying changes in swimming performance, the contractile properties of fast muscle fibres were determined in fish of similar body length. The rate of tetanic force relaxation (time from last stimulus to 50% peak force) was 34% faster in gr avid than in postspawning fish. Maximum contraction velocity, determined by the slack-test method, was significantly higher in gravid than in postspawning fish (6·8 v . 5·9 muscle lengths s⁻¹). In contrast, both maximum isometric stress and power output (determined from the force–velocity relationship) were >50% higher in fibres from postspawning than from gravid fish, even though myofibrillar protein and water contents were similar (120 mg g⁻¹ wet mass and 86% of body mass, respectively). The results show that swimming performance and the contractile properties of fast muscle fibres vary with the reproductive cycle in short-horn sculpin acclimated to the same photoperiodic and temperature regime.