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.     

How small puffers (Teleostei: Tetraodontidae) swim

 The tetraodontiform swimming mode has recently attracted attention because puffers swim very steadily and, unlike most of the other median and paired fin (MPF) swimmers, use more than one pair of fins to propel themselves through the water. To date, only one study presenting data concerning the swimming kinematics of puffers has been published, and this study dealt only with two species of large body size. In the present study, the swimming kinematics of small puffers (<6 cm TL) Tetraodon schoutedeni is described and compared to the swimming kinematics of larger puffers and boxfish. The results show that, generally, the swimming kinematics of small puffers is similar to that of larger puffers. The main differences that were found are in the synchronization of dorsal and anal fin motion, and in the motion of the pectoral fins, which complete their adduction before the dorsal and anal fins do. Maximum fin beat frequency was 18.4 Hz, much faster than that of larger puffers. At slow and median swimming speeds, dorsal fin beat amplitude increases with swimming speed and then remains constant between median and fast swimming speeds. The results confirm previous findings that puffers swim extremely steadily. Most of the differences in swimming kinematics between large and small puffers can be attributed to the size differences, but the difference in fin synchronization should be further studied to be completely understood. Received: September 27, 2002 / Revised: January 7, 2003 / Accepted: February 6, 2003     

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.     

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.     

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.     

实验室条件下唐鱼两性异形及其与游泳能力关系

为了解唐鱼两性异形及其与游泳能力关系,检测了性成熟阶段唐鱼躯干部和鱼鳍形态特征以及爆发游泳速度(U_burst)和临界游泳速度(U_crit)在雌雄之间的差异,旨在从形态适应角度探究长期进化中雌雄唐鱼各自面对选择压力所产生的游泳能力差异及其机制,从而为野生唐鱼保护提供基础数据.结果表明: 雌性唐鱼的体长、头高、头宽、尾鳍面积以及吻端至枕骨后末端、腹鳍起点至背鳍末端等长度均与雄性无显著差异.而体高、体宽、腹鳍起点至背鳍起点等反映腹腔大小的形态参数以及吻端至背鳍起点、吻端至臀鳍起点、枕骨后末端至背鳍起点等反映躯干部大小的形态参数均显示为雌性显著大于雄性,但头长以及胸鳍面积、腹鳍面积、背鳍面积和臀鳍面积均显示为雄性显著大于雌性.对所有数据进行主成分分析,结果显示第1主成分贡献率为74.2%,负载量较大的是体长、头长、头高、体高、头宽、体宽以及各鳍之间距离等主要反映唐鱼躯干整体特征的参数;第2主成分贡献率为15.7%,负载量较大的是胸鳍面积、腹鳍面积、背鳍面积和臀鳍面积等主要反映鱼鳍特征的参数.唐鱼性别在第1主成分上无法区分,但在第2主成分却可以明显区分.根据体宽、胸鳍面积、腹鳍面积、背鳍面积和臀鳍面积等建立的性别判别方程对雌雄判断准确率达到91.8%～92.5%.唐鱼游泳能力测定结果显示,雌性U_burst与雄性无显著差异,但U_crit显著小于雄性.以上结果表明,雌雄唐鱼两性异形主要集中在与游泳能力相关的鱼鳍特征上.相比雄性,雌性唐鱼虽然胸鳍等鱼鳍面积较小导致其U_crit小于雄性,却具有更长的躯干部以保证其同样具有较高的爆发游泳能力,从而有利于在流速波动很大的溪流中躲避捕食和进行其他应急活动;相比雌性,雄性唐鱼则具有较大的鱼鳍面积保证其U_crit高于雌性,以利于日常活动及在繁殖过程中追逐雌性等相对持久性游泳运动.     

Bluegill Lepomis macrochirus synchronize pectoral fin motion and opercular pumping

The relative timing between operculum and pectoral fin motion was examined in swimming bluegill Lepomis macrochirus to determine if respiratory fluid flows from the operculum might have an effect on flow over the pectoral fin. Five bluegill were filmed swimming at speeds from 0·5 to 1·5 body (total) lengths s⁻¹. The timing of opercular pumping and pectoral fin beating was noted and analysed using circular statistics. Fish tended to ventilate their gills every second or third pectoral fin beat. While locomotion and ventilation had different frequencies, however, they were synchronized: fish maintained a consistent phase relationship between them. Thus, within pectoral fin beats when the operculum pumps, the jet consistently occurred during pectoral fin abduction, ending just after the fin was fully abducted and beginning adduction. Based on the distance between the opercular slit and the pectoral fin base, the jet was estimated to reach the fin during maximum abduction. Dye flow visualization confirmed this estimate, revealing that the opercular flow wraps around the base of the fin during peak abduction, when it is likely to have little hydrodynamic effect.     

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.     

Gait transition speed, pectoral fin-beat frequency and amplitude in Cymatogaster aggregata, Embiotoca lateralis and Damalichthys vacca

Surfperches are labriform swimmers and swim primarily with their pectoral fins, using the tail to assist only at higher speeds. The transition, from pectoral to pectoral and caudal fins, occurs at a threshold speed that has been termed physiologically and biomechanically 'equivalent' for fishes of different size. The gait transition ( U _P-C) of Cymatogaster aggregata occurred at a higher speed (measured in bodylengths s⁻¹) for smaller fish than larger fish. At U _P-C, pectoral fin-beat frequency was size-dependent: smaller fish have a higher pectoral fin-beat frequency than larger fish. In contrast, at low speeds (i.e. <60% of U _P-C) the pectoral fin-beat frequency was independent of the size of the fish. Inter-specific comparisons of U _P-C, pectoral fin-beat frequency and amplitude among C. aggregata, Embiotoca lateralis and Damalichthys vacca showed that C. aggregata had a higher U _P-C than E. lateralis and D. vacca . The pectoral fin-beat frequency at U _P-C showed no significant differences among species. Cymatogaster aggregata achieved higher U _P-C, in part, through increased fin beat amplitude rather than frequency. These differences in performance may be related to the different habitats in which these species live.     

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.     

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.     

Kinematics of swimming of penguins at the Detroit Zoo

Kinematic parameters were examined in a study of the swimming abilities of seven species of penguins housed at the Detroit Zoo. Penguins produce thrust over both halves of the wing stroke cycle, as observed in fishes using the caudal or pectoral fins for locomotion, but not in other birds in level forward flight. Unpowered gliding phases between wing strokes were observed in all species at swimming speeds less than 1.25 m/sec, while Emperor, King and Adelie penguins interpose gliding phases over a broad range of speeds. Videotape records reveal that length-specific speed is correlated with increases in wingbeat frequency and, for most of the species examined, stride length. These findings are in contrast to those reported for other, flying birds, which maintain a relatively constant wingbeat frequency but vary stride length with forward speed, and for most fishes, which vary speed with tailbeat frequency but maintain a constant stride length. The results are somewhat comparable to those reported for Cymatogaster , a fish which uses the pectoral fins for locomotion. Drag coefficients of three gliding Emperor penguins were 2.1, 3.0 and 3.0 × 10-⁻³ at Reynolds numbers of 1.25, 1.62 and 1.76 × 10⁶, respectively.     

A new approach to quantifying morphological variation in bluegill Lepomis macrochirus

Bluegill Lepomis macrochirus showed intraspecific morphological and behavioural differences dependent on the environment. Pelagic L. macrochirus had more fusiform bodies, a higher pectoral fin aspect ratio, a larger spiny dorsal fin area and pectoral fins located farther from the centre of mass than littoral L. macrochirus (P < 0·05). The shape of the body and pectoral fins, in particular, were suggestive of adaptation for sustained high-speed and economical labriform swimming. Littoral L. macrochirus had a deeper and wider body, deeper caudal fins and wider mouths than pelagic L. macrochirus (P < 0·05). Additionally, the soft dorsal, pelvic, anal and caudal fins of littoral L. macrochirus were positioned farther from the centre of mass (P < 0·05). The size and placement of these fins suggested that they will be effective in creating turning moments to facilitate manoeuvring in the macrophyte-dense littoral habitat.     

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.     

The locomotory function of the fins in the squid Loligo pealei

Kinematic data of high spatial and temporal resolution, acquired from image sequences of adult long-finned squid, Loligo pealei, during steady swimming in a flume, were used to examine the role of fins and the coordination between fin and jet propulsion in squid locomotion. Fin shape and body outlines were digitized and used to calculate fin wave speed, amplitude, frequency, angle of attack, body deformation, speed, and acceleration. L. pealei were observed to have two fin gait patterns with a transition at 1.4-1.8 mantle lengths per second (L_m s^-1) marked by alternation between the two patterns. Fin motion in L. pealei exhibited characteristics of both traveling waves and flapping wings. At low speeds, fin motion was more wave-like; at high speeds, fin motion was more flap-like and was marked by regular periods during which the fins were wrapped tightly against the mantle. Fin cycle frequencies were dependent on swimming speed and gait, and obvious coordination between the fins and jet were observed. Fin wave speed, angle of attack, and body acceleration confirmed the role of fins in thrust production and revealed a role of fins at all swimming speeds by a transition from drag-based to lift-based thrust when fin wave speed dropped below swimming speed. Estimates of peak fin thrust were as high as 0.44-0.96 times peak jet thrust in steady swimming over the range of swimming speeds observed. Fin downstrokes generally contributed more to thrust than did upstrokes, especially at high speeds.     

The role of the pectoral fins in body trim of sharks

In a large aquarium the leopard shark Triakis semifasciata , sand tiger shark Odontaspis taurus , sandbar shark Carcharhinus plumbeus , and spiny dogfish Squalus acanthias cruised steadily at 0·1-0·7 body lengths s^-1. Relative to the trajectory of the shark, the pectoral fins were maintained at a positive angle of ttack regardless of vertical direction. For level swimming the mean angle of attack for the pectoral fin was 11±1·7, 10·1±1·3°, 9·3±1·3°, and 15·0±0·0 for T. semifasciata , C. plumbeus , O. taurus , and S. acanthias , respectively. The long axis of the body was canted at an angle of attack for T. semifasciata and S. acanthias , but trim was maintained during level swimming for C. plumbeus and O. taurus . Hydrodynamic analysis of the body and fin design of T. semifasciata indicated that the pectoral fins could develop suffcient pitching moment to maintain depth and keep the body in trim. Demonstration of positive angles of attack support the hypothesis that lift is generated in the anterior body to counterbalance the lift produced by the heterocercal tail.     

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.     

Larval development of Argentine hake Merluccius hubbsi

The ontogeny of the developmental stages of the hake Merluccius hubbsi is described. Fish larvae and post-transitional juveniles were collected in the Nor-Patagonian area from 1989 to 2004. The opening of the mouth and the pigmentation of the eyes are coincident with yolk resorption, finishing the yolk-sac stage. This species presents pigmentation on the head, trunk and tail typical of gadiform larvae. Pectoral fin development is completed during the transformation stage. The post-transitional juvenile stage begins when the fin-ray complements are complete and squamation begins. The fins become fully formed in the following sequence: pelvic fins, first dorsal fin, second dorsal and anal fins together, caudal fin and pectoral fins. The caudal complex is totally developed in larvae of 22·0–23·0 mm standard lengths ( L _S) and all vertebral elements are first observed in larvae of 8·5 mm L _S. The rate of development of M. hubbsi observed in this study could be faster than the rates reported for other species of Merluccius by different authors.     

Functional Morphology of Aquatic Flight in Fishes: Kinematics, Electromyography, and Mechanical Modeling of Labriform Locomotion

Labriform locomotion is the primary swimming mode for many fishesthat use the pectoral fins to generate thrust across a broadrange of speeds. A review of the literature on hydrodynamics,kinematics, and morphology of pectoral fin mechanisms in fishesreveals that we lack several kinds of morphological and kinematicdata that are critical for understanding thrust generation inthis mode, particularly at higher velocities. Several needsinclude detailed three-dimensional kinematic data on speciesthat are pectoral fin swimmers across a broad range of speeds,data on the motor patterns of pectoral fin muscles, and thedevelopment of a mechanical model of pectoral fin functionalmorphology. New data are presented here on pectoral fin locomotionin Gomphosus varius, a labrid fish that uses the pectoral finsat speeds of 1 –6 total body lengths per second. Three-dimensionalkinematic data for the pectoral fins of G. varius show thata typical "drag-based" mechanism is not used in this species.Instead, the thrust mechanics of this fish are dominated bylift forces and acceleration reaction forces. The fin is twistedlike a propeller during the fin stroke, so that angles of attackare variable along the fin length. Electromyographic data onsix fin muscles indicate the sequence of muscle activity thatproduces antagonistic fin abduction and adduction and controlsthe leading edge of the fin. EMG activity in abductors and adductorsis synchronous with the start of abduction and adduction, respectively,so that muscle mechanics actuate the fin with positive work.A mechanical model of the pectoral fin is proposed in whichfin morphometrics and computer simulations allow predictionsof fin kinematics in three dimensions. The transmission of forceand motion to the leading edge of the fin depends on the mechanicaladvantage of fin ray levers. An integrative program of researchis suggested that will synthesize data on morphology, physiology,kinematics, and hydrodynamics to understand the mechanics ofpectoral fin swimming.     

Development of the pectoral, pelvic, dorsal and anal fins in cultured sea bream

The pectoral fin girdle was the first element of the fins to develop in Sparus aurata. By 3·1mm L _N (notochord length) the cleithrum was ossified and the cartilaginous caracoid-scapula was present. The fin was fully developed at 11·6 mm L _S (standard length) and by 16·0 mm L _S most elements of the fin were ossified. The pelvic fins were the last pair to develop and rudiments of these were first detected at 7·9 mm L _S. The pelvic fin and girdle were completely formed and ossified at 16·0 mm L _S. The development of dorsal and anal fins began at c. 6·5–7·0 mm L _S with the formation of 10 cartilaginous dorsal proximal radials and eight cartilaginous ventral proximal radials. The three cartilaginous predorsals (supraneurals) appeared at 7·7 mm L _S and the ossification of dorsal and anal proximal and distal radials began, respectively, at 10·5 mm L _S and 11·3 mm L _S. Ossified structures in the fins were also classified according to their origin, as being either dermal or endochondral. Finally the chronology of appearance of fin structures in S. aurata was compared with that reported for other Sparidae, Engraulidae and Haemulidae.