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Activation of Erk and JNK MAPK pathways by acute swim stress in rat brain regions.

BACKGROUND: The mitogen-activated protein kinases (MAPKs) have been shown to participate in a wide array of cellular functions. A role for some MAPKs (e.g., extracellular signal-regulated kinase, Erk1/2) has been documented in response to certain physiological stimuli, such as ischemia, visceral pain and electroconvulsive shock. We recently demonstrated that restraint stress activates the Erk MAPK pathway, but not c-Jun-N-terminal kinase/stress-activated protein kinase (JNK/SAPK) or p38MAPK, in several rat brain regions. In the present study, we investigated the effects of a different stressor, acute forced swim stress, on the phosphorylation (P) state of these MAPKs in the hippocampus, neocortex, prefrontal cortex, amygdala and striatum. In addition, effects on the phosphorylation state of the upstream activators of the MAPKs, their respective MAPK kinases (MAPKKs; P-MEK1/2, P-MKK4 and P-MKK3/6), were determined. Finally, because the Erk pathway can activate c-AMP response element (CRE) binding (CREB) protein, and swim stress has recently been reported to enhance CREB phosphorylation, changes in P-CREB were also examined. RESULTS: A single 15 min session of forced swimming increased P-Erk2 levels 2-3-fold in the neocortex, prefrontal cortex and striatum, but not in the hippocampus or amygdala. P-JNK levels (P-JNK1 and/or P-JNK2/3) were increased in all brain regions about 2-5-fold, whereas P-p38MAPK levels remained essentially unchanged. Surprisingly, levels of the phosphorylated MAPKKs, P-MEK1/2 and P-MKK4 (activators of the Erk and JNK pathways, respectively) were increased in all five brain regions, and much more dramatically (P-MEK1/2, 4.5 to > 100-fold; P-MKK4, 12 to approximately 300-fold). Consistent with the lack of forced swim on phosphorylation of p38MAPK, there appeared to be no change in levels of its activator, P-MKK3/6. P-CREB was increased in all but cortical (prefrontal, neocortex) areas. CONCLUSIONS: Swim stress specifically and markedly enhanced the phosphorylation of the MAPKKs P-MEK1/2 and P-MKK4 in all brain regions tested without apparent alteration in the phosphorylation of P-MKK3/6. Curiously, phosphorylation of their cognate substrates (Erk and JNK) was increased to a much more modest extent, and in some brain regions was not altered. Similarly, there was a region-specific discrepancy between Erk and CREB phosphorylation. Possible explanations for these findings and comparison with the effects of restraint stress will be discussed.

Animals↗

Condition, prolonged swimming performance and muscle metabolic capacities of cod Gadus morhua.

This study evaluated the link between swimming endurance and condition of Atlantic cod Gadus morhua that had been fed or starved during the 16 weeks preceding the tests, and assessed whether muscle metabolic capacities explain such links. The condition factor [(somatic mass x fork length(-3))x100] of starved cod was 0.54+/-0.1 whereas that of fed cod was 0.81+/-0.1. In white and red muscle, we measured four glycolytic enzymes: phosphofructokinase (PFK), pyruvate kinase (PK), creatine kinase (CK) and lactate dehydrogenase (LDH), two mitochondrial enzymes: cytochrome c oxidase (CCO) and citrate synthase (CS), a biosynthetic enzyme, nucleoside diphosphate kinase (NDPK), glycogen and protein levels and water content. Muscle samples were taken at three positions along the length of the fish; starvation affected the metabolic capacities of white muscle more than those of red muscle. The levels of glycolytic enzymes and glycogen changed more in white than red muscle during starvation. Both in fed and starved cod, muscle metabolic capacities varied with position along the fish; starvation reduced this longitudinal variation more in white than red muscle. In white muscle of fed cod, the glycolytic enzyme levels increased from head to tail, while in starved cod this longitudinal variation disappeared. In red muscle mitochondrial enzyme levels were highest in the caudal sample, but fewer differences were found for glycolytic enzymes. Swimming endurance was markedly affected by fish condition, with starved fish swimming only 30% of the time (and distance) of fed fish. This endurance was closely linked with the number of burst-coast movements during the test and the activity of CCO and LDH in white muscle. The number of burst-coast movements was significantly linked with condition factor and PFK activity in caudal red muscle and gill arch mass. Our data indicated that cod use both glycolytic and oxidative capacities to support endurance swimming. Furthermore, swimming endurance is linked with the metabolic capacities of red and white muscle.

Animals↗

The effect of temperature on swimming performance and oxygen consumption in adult sockeye (Oncorhynchus nerka) and coho (O. kisutch) salmon stocks.

Our knowledge of the swimming capabilities and metabolic rates of adult salmon, and particularly the influence of temperature on them, is extremely limited, and yet this information is critical to understanding the remarkable upstream migrations that these fish can make. To remedy this situation, we examined the effects of temperature on swimming performance and metabolic rates of 107 adult fish taken from three stocks of sockeye salmon Oncorhynchus nerka and one stock of coho salmon O. kisutch at various field and laboratory locations, using large, portable, swim tunnels. The salmon stocks were selected because of differences in their ambient water temperature (ranging from 5 degrees C to 20 degrees C) and the total distance of their in-river migrations (ranging from approximately 100 km for coastal stocks to approximately 1100 km for interior stocks). As anticipated, differences in routine metabolic rate observed among salmon stocks were largely explained by an exponential dependence on ambient water temperature. However, the relationship between water temperature and maximum oxygen consumption (MO2max), i.e. the MO2 measured at the critical swimming speed (Ucrit), revealed temperature optima for MO2max that were stock-specific. These temperature optima were very similar to the average ambient water temperatures for the natal stream of a given stock. Furthermore, at a comparable water temperature, the salmon stocks that experienced a long and energetically costly in-river migration were characterized by a higher MO2max, a higher scope for activity, a higher Ucrit and, in some cases, a higher cost of transport, relative to the coastal salmon stocks that experience a short in-river migration. We conclude that high-caliber respirometry can be performed in a field setting and that stock-specific differences in swimming performance of adult salmon may be important for understanding upstream migration energetics and abilities.

Animal Migration↗

Excess post-exercise oxygen consumption in adult sockeye (Oncorhynchus nerka) and coho (O. kisutch) salmon following critical speed swimming.

The present study measured the excess post-exercise oxygen cost (EPOC) following tests at critical swimming speed (Ucrit) in three stocks of adult, wild, Pacific salmon (Oncorhynchus sp.) and used EPOC to estimate the time required to return to their routine level of oxygen consumption (recovery time) and the total oxygen cost of swimming to Ucrit. Following exhaustion at Ucrit, recovery time was 42-78 min, depending upon the fish stock. The recovery times are several-fold shorter than previously reported for juvenile, hatchery-raised salmonids. EPOC varied fivefold among the fish stocks, being greatest for Gates Creek sockeye salmon (O. nerka), which was the salmon stock that had the longest in-river migration, experienced the warmest temperature and achieved the highest maximum oxygen consumption compared with the other salmon stocks that were studied. EPOC was related to Ucrit, which in turn was directly influenced by ambient test temperature. The non-aerobic cost of swimming to Ucrit was estimated to add an additional 21.4-50.5% to the oxygen consumption measured at Ucrit. While these non-aerobic contributions to swimming did not affect the minimum cost of transport, they were up to three times higher than the value used previously for an energetic model of salmon migration in the Fraser River, BC, Canada. As such, the underestimate of non-aerobic swimming costs may require a reevaluation of the importance of how in-river barriers like rapids and bypass facilities at dams, and year-to-year changes in river flows and temperatures, affect energy use and hence migration success.

Animal Migration↗

How do cormorants counter buoyancy during submerged swimming?

Buoyancy is a de-stabilizing force for diving cormorants that forage at shallow depths. Having to counter this force increases the cost of transport underwater. Cormorants are known to be less buoyant than most water birds but are still highly buoyant (rho= approximately 0.8 kg m(-3)) due to their adaptations for aerial flight. Nevertheless, cormorants are known to dive at a wide range of depths, including shallow dives where buoyancy is maximal. We analyzed the kinematics of underwater swimming of the great cormorant (Phalacrocorax carbo sinensis) in a shallow pool to discover and evaluate the mechanisms countering buoyancy while swimming horizontally. The birds maintained a very uniform cyclic paddling pattern. Throughout this cycle, synchronized tilting of the body, controlled by the tail, resulted in only slight vertical drifts of the center of mass around the average swimming path. We suggest that this tilting behavior serves two purposes: (1) the elongated bodies and the long tails of cormorants, tilted at a negative angle of attack relative to the swimming direction, generate downward directed hydrodynamic lift to resist buoyancy and (2) during the propulsive phase, the motion of the feet has a significant vertical component, generating a vertical component of thrust downward, which further helps to offset buoyancy. The added cost of the drag resulting from this tilting behavior may be reduced by the fact that the birds use a burst-and-glide pattern while swimming.

Animals↗

Partitioning heat loss from mallard ducklings swimming on the air-water interface.

Water birds whose young begin swimming while downy are interesting because hypothermia and mortality are associated with wetting. While wetting is known to increase heat loss, little is known about basic issues, such as the amount of heat lost to air vs water during surface swimming. To partition heat loss to air and water, we measured the body temperature, metabolism and thermal conductance of 2-3-day-old mallard ducklings (Anas platyrhynchos) swimming under different combinations of air and water temperature. Ventral down remained dry or was wetted only on the surface, and most ducklings could maintain Tb>39 degrees C for 1 h while swimming on water as cold as 5 degrees C. Ducklings were at or below thermal neutrality when swimming in water at Tw=30 degrees C even when air temperature Ta=45 degrees C. Heat loss from ducklings with dry down to air and water was partitioned by fitting data to a heat transfer model of the form M=G(Tb-Tw)+Ke(Tb-Ta). For an average 48 g duckling, thermal conductance to water increased with water temperature, G=0.0470(1+1.059 x 10(-6)Tw4)W/ degrees C-animal. Conductance to air was Ke=0.0196 W/ degrees C-animal for all air temperatures. Thus, a minimum of 70% of metabolic heat production is lost to water, and this fraction increases with increasing temperature.

Animals↗

Cardiac preload and venous return in swimming sea bass (Dicentrarchus labrax L.).

Cardiac preload (central venous pressure, P(CV), mean circulatory filling pressure (MCFP), dorsal aortic blood pressure (P(DA)) and relative cardiac output (Q) were measured in sea bass (Dicentrarchus labrax) at rest and while swimming at 1 and 2 BL s(-1). MCFP, an index of venous capacitance and the upstream venous pressure driving the return of venous blood to the heart, was measured as the plateau in Pcv during ventral aortic occlusion. Compared with resting values, swimming at 1 and 2 BL s(-1) increased Q (by 15+/-1.5 and 38+/-6.5%, respectively), P(CV) (from 0.11+/-0.01 kPa to 0.12+/-0.01 and 0.16+/-0.02 kPa, respectively), MCFP (from 0.27+/-0.02 kPa to 0.31+/-0.02 and 0.40+/-0.04 kPa, respectively) and the calculated pressure gradient for venous return (DeltaP(V), from 0.16+/-0.01 kPa to 0.18+/-0.02 and 0.24+/-0.02 kPa, respectively), but not P(DA). In spite of an increased preload, the increase in Q was exclusively mediated by an increased heart rate (f(H), from 80+/-4 beats min(-1) to 88+/-4 and 103+/-3 beats min(-1), respectively), and stroke volume (Vs) remained unchanged. Prazosin treatment (1 mg kg(-1) Mb) abolished pressure and flow changes during swimming at 1 BL s(-1), but not 2 BL s(-1), indicating that other control systems besides an alpha-adrenoceptor control are involved. This study is the first to address the control of venous capacitance in swimming fish. It questions the generality that increased Q during swimming is regulated primarily through Vs and shows that an increased cardiac filling pressure does not necessarily lead to an increased Vs in fish, but may instead compensate for a reduced cardiac filling time.

Animals↗

Emperor penguins adjust swim speed according to the above-water height of ice holes through which they exit.

Emperor penguins leap from the water onto the sea ice. Their ability to reach above-water height depends critically on initial vertical speed of their leaping, assuming that the kinetic energy is converted to gravitational potential energy. We deliberately changed the above-water heights of ice hole exits, in order to examine whether penguins adjusted swim speed in accordance with the above-water height of the ice. Penguins were maintained in a corral on the fast ice in Antarctica, and voluntarily dived through two artificial ice holes. Data loggers were deployed on the penguins to monitor under water behavior. Nine instrumented penguins performed 386 leaps from the holes during experiments. The maximum swim speeds within 1 s before the exits through the holes correlated significantly with the above-water height of the holes. Penguins adopted higher speed to exit through the higher holes than through the lower holes. Speeds of some failed exits were lower than the theoretical minimum values to reach a given height. Penguins failed to exit onto the sea ice in a total of 37 of the trials. There was no preference to use lower holes after they failed to exit through the higher holes. Rather, swim speed was increased for subsequent attempts after failed leaps. These data demonstrated that penguins apparently recognized the above-water height of holes and adopted speeds greater than the minimal vertical speeds to reach the exit height. It is likely, especially in the case of higher holes (>40 cm), that they chose minimum speeds to exit through the holes to avoid excess energy for swimming before leaping. However, some exceptionally high speeds were recorded when they directly exited onto the ice from lower depths. In those cases, birds could increase swim speed without strokes for the final seconds before exit and they only increased the steepness of their body angles as they surfaced, which indicates that the speed required for leaps by emperor penguins were aided by buoyancy, and that penguins can sometimes exit through the ice holes without any stroking effort before leaping.

Animals↗

On the estimation of swimming and flying forces from wake measurements.

The transfer of momentum from an animal to fluid in its wake is fundamental to many swimming and flying modes of locomotion. Hence, properties of the wake are commonly studied in experiments to infer the magnitude and direction of locomotive forces. The determination of which wake properties are necessary and sufficient to empirically deduce swimming and flying forces is currently made ad hoc. This paper systematically addresses the question of the minimum number of wake properties whose combination is sufficient to determine swimming and flying forces from wake measurements. In particular, it is confirmed that the spatial velocity distribution (i.e. the velocity field) in the wake is by itself insufficient to determine swimming and flying forces, and must be combined with the fluid pressure distribution. Importantly, it is also shown that the spatial distribution of rotation and shear (i.e. the vorticity field) in the wake is by itself insufficient to determine swimming and flying forces, and must be combined with a parameter that is analogous to the fluid pressure. The measurement of this parameter in the wake is shown to be identical to a calculation of the added-mass contribution from fluid surrounding vortices in the wake, and proceeds identically to a measurement of the added-mass traditionally associated with fluid surrounding solid bodies. It is demonstrated that the velocity/pressure perspective is equivalent to the vorticity/vortex-added-mass approach in the equations of motion. A model is developed to approximate the contribution of wake vortex added-mass to locomotive forces, given a combination of velocity and vorticity field measurements in the wake. A dimensionless parameter, the wake vortex ratio (denoted Wa), is introduced to predict the types of wake flows for which the contribution of forces due to wake vortex added-mass will become non-negligible. Previous wake analyses are re-examined in light of this parameter to infer the existence and importance of wake vortex added-mass in those cases. In the process, it is demonstrated that the commonly used time-averaged force estimates based on wake measurements are not sufficient to prove that an animal is generating the locomotive forces necessary to sustain flight or maintain neutral buoyancy.

Animals↗

Unifying constructal theory for scale effects in running, swimming and flying.

Biologists have treated the view that fundamental differences exist between running, flying and swimming as evident, because the forms of locomotion and the animals are so different: limbs and wings vs body undulations, neutrally buoyant vs weighted bodies, etc. Here we show that all forms of locomotion can be described by a single physics theory. The theory is an invocation of the principle that flow systems evolve in such a way that they destroy minimum useful energy (exergy, food). This optimization approach delivers in surprisingly direct fashion the observed relations between speed and body mass (M(b)) raised to 1/6, and between frequency (stride, flapping) and M(b)(-1/6), and shows why these relations hold for running, flying and swimming. Animal locomotion is an optimized two-step intermittency: an optimal balance is achieved between the vertical loss of useful energy (lifting the body weight, which later drops), and the horizontal loss caused by friction against the surrounding medium. The theory predicts additional features of animal design: the Strouhal number constant, which holds for running as well as flying and swimming, the proportionality between force output and mass in animal motors, and the fact that undulating swimming and flapping flight occur only if the body Reynolds number exceeds approximately 30. This theory, and the general body of work known as constructal theory, together now show that animal movement (running, flying, swimming) and fluid eddy movement (turbulent structure) are both forms of optimized intermittent movement.

Animals↗

Regulation of stroke pattern and swim speed across a range of current velocities: diving by common eiders wintering in polynyas in the Canadian Arctic.

Swim speed during diving has important energetic consequences. Not only do costs increase as drag rises non-linearly with increasing speed, but speed also affects travel time to foraging patches and therefore time and energy budgets over the entire dive cycle. However, diving behaviour has rarely been considered in relation to current velocity. Strong tidal currents around the Belcher Islands, Nunavut, Canada, produce polynyas, persistent areas of open water in the sea ice which are important habitats for wildlife wintering in Hudson Bay. Some populations of common eiders Somateria mollissima sedentaria remain in polynyas through the winter where they dive to forage on benthic invertebrates. Strong tidal currents keep polynyas from freezing, but current velocity can exceed 1.5 m s(-1) and could influence time and energy costs of diving and foraging. Polynyas therefore provide naturally occurring flume tanks allowing investigation of diving strategies of free ranging birds in relation to current velocity. We used a custom designed sub-sea ice camera to non-invasively investigate over 150 dives to a depth of 11.3 m by a population of approximately 100 common eiders at Ulutsatuq polynya during February and March of 2002 and 2003. Current speed during recorded dives ranged from 0 to 1 m s(-1). As currents increased, vertical descent speed of eiders decreased, while descent duration and the number of wing strokes and foot strokes during descent to the bottom increased. However, nearly simultaneous strokes of wings and feet, and swim speed relative to the moving water, were maintained within a narrow range (2.28+/-0.23 Hz; 1.25+/-0.14 m s(-1), respectively). This close regulation of swim speed over a range in current speed of 1.0 m s(-1) might correspond to efficient muscle contraction rates, and probably reduces work rates by avoiding rapidly increasing drag at greater speeds; however, it also increases travel time to benthic foraging patches. Despite regulation of average swim speed, high instantaneous speeds during oscillatory stroking can increase dive costs due to drag. While most diving birds have been considered either foot or wing propelled, eider ducks used both wing and foot propulsion during descent. Our observations indicate that the power phase of foot strokes coincides with the transition between upstroke and downstroke of the wings, when drag is greatest. Coordinated timing between foot and wing propulsion could therefore serve to maintain a steadier speed during descent and decrease the costs of diving. Despite tight regulation of stroke and swim speed patterns, descent duration and total number of foot and wing strokes during descent increase non-linearly with increasing current velocity, suggesting an increase in energetic costs of diving.

Animals↗

Swimming in the California sea lion: morphometrics, drag and energetics.

During swimming, the California sea lion, Zalophus californianus (Lesson), generates thrust forces solely by means of its pectoral flippers. This study examines the drag, energetic cost and efficiency associated with this method of locomotion. Sea lions are highly streamlined, with a fineness ratio of 5.5 and maximum girth at 40% of body length. This profile leads to reduced drag and swimming power requirements. Films of gliding animals showed the drag coefficient (based on wetted surface area) to be 0.0042 at a Reynolds number of 2.0 X 10(6). This value is comparable to that found for other aquatic vertebrates and suggests that the sea lion's morphology helps to delay turbulent separation and maintain laminar flow over the forward portion of its body. Swimming metabolism was measured in a water flume at velocities up to 1.3 ms-1. Effective swimming speeds up to 2.7 ms-1 were attained by increasing each animal's drag. Oxygen consumption rose exponentially with velocity and for two animals was best described as VO2 = 6.27e0.48U, where VO2 is in mlO2 min-1 kg-1 and U is in ms-1. Minimum cost of transport for these animals was 0.12 ml O2 kg-1 m-1 at a relative speed of 1.4 body lengths s-1. This is 2.5 times that predicted for a fish of similar size. Swimming efficiencies were determined from these results using power output values calculated from the measured drag coefficient and standard hydrodynamic equations. At the highest velocity, aerobic efficiency reached a maximum of 15% while mechanical efficiency of the foreflippers was 80%. The results demonstrate that foreflipper propulsion is a highly efficient and comparatively inexpensive method of locomotion in aquatic mammals.

Animals↗

Heart rate and stroke volume contribution to cardiac output in swimming yellowfin tuna: response to exercise and temperature.

Cardiac performance in the yellowfin tuna (Thunnus albacares, 673-2470 g, 33-53 cm fork length, FL) was examined in unanesthetized fish swimming in a large water tunnel. Yellowfin tuna were fitted with either electrocardiogram electrodes or a transcutaneous Doppler blood-flow probe over the ventral aorta and exposed to changes in swimming velocity (range 0.8-2.9 FLs-1) or to an acute change in temperature (18-28 degrees C). Heart rates (fH) at +/-1 degree C (30-130 beats min-1) were lower on average than previous measurements with non-swimming (restrained) tunas and comparable with those for other active teleosts at similar relative swimming velocities. Although highly variable among individuals, fH increased with velocity (U, in FLs-1) in all fish (fH = 17.93U + 49.93, r2 = 0.14, P < 0.0001). Heart rate was rapidly and strongly affected by temperature (Q10 = 2.37). Blood flow measurements revealed a mean increase in relative cardiac output of 13.6 +/- 3.0% with exercise (mean velocities 1.23-2.10 FLs-1) caused by an 18.8 +/- 5.4% increase in fH and a 3.9 +/- 2.3% decrease in stroke volume. These results indicate that, unlike most other fishes, cardiac output in yellowfin tuna is regulated primarily through increases in fH. Acute reductions in ambient temperature at slow swimming velocities resulted in decreases in cardiac output (Q10 = 1.52) and fH (Q10 = 2.16), but increases in stroke volume (Q10 = 0.78). This observation suggests that the lack of an increase in stroke volume during exercise is not due to the tuna heart operating at maximal anatomical limits.

Animals↗

Diving and swimming performance of white whales, Delphinapterus leucas: an assessment of plasma lactate and blood gas levels and respiratory rates.

The white whale Delphinapterus leucas is an exceptional diver, yet we know little about the physiology that enables this species to make prolonged dives. We studied trained white whales with the specific goal of assessing their diving and swimming performance. Two adult whales performed dives to a test platform suspended at depths of 5-300 m. Behavior was monitored for 457 dives with durations of 2.2-13.3 min. Descent rates were generally less than 2 m s-1 and ascent rates averaged 2.2-3 m s-1. Post-dive plasma lactate concentration increased to as much as 3.4 mmol l-1 (4-5 times the resting level) after dives of 11 min. Mixed venous PO2 measured during voluntary breath-holds decreased from 79 to 20 mmHg within 10 min; however, maximum breath-hold duration was 17 min. Swimming performance was examined by training the whales to follow a boat at speeds of 1.4-4.2 m s-1. Respiratory rates ranged from 1.6 breaths min-1 at rest to 5.5 breaths min-1 during exercise and decreased with increasing swim speed. Post-exercise plasma lactate level increased to 1.8 mmol l-1 (2-3 times the resting level) following 10 min exercise sessions at swimming speeds of 2.5-2.8 m s-1. The results of this study are consistent with the calculated aerobic dive limit (O2 store/metabolic rate) of 9-10 min. In addition, white whales are not well adapted for high-speed swimming compared with other small cetaceans.

Animals↗

Fish swimming: patterns in muscle function.

Undulatory swimming in fish is powered by the segmental body musculature of the myotomes. Power generated by this muscle and the interactions between the fish and the water generate a backward-travelling wave of lateral displacement of the body and caudal fin. The body and tail push against the water, generating forward thrust. The muscle activation and strain patterns that underlie body bending and thrust generation have been described for a number of species and show considerable variation. This suggests that muscle function may also vary among species. This variation must be due in large part to the complex interactions between muscle mechanical properties, fish body form, swimming mode, swimming speed and phylogenetic relationships. Recent work in several laboratories has been directed at studying patterns of muscle power output in vitro under simulated swimming conditions. This work suggests that the way that fish generate muscle power and convert it into thrust through the body and caudal fin does indeed vary. However, despite the differences, several features appear to be common to virtually all species studied and suggest where future effort should be directed if muscle function in swimming fish is to be better understood.

Animals↗

Power production during steady swimming in largemouth bass and rainbow trout.

Steady swimming in fishes is powered by the aerobic or red muscle, but there are conflicting theories on the relative roles of the anterior and posterior red muscle in powering steady swimming. To examine how red muscle is used to power steady swimming in rainbow trout (Oncorhynchus mykiss), electromyographic (EMG) and sonomicrometry recordings were made of muscle activity in vivo. These data were used in in vitro work-loop studies of muscle power production. Data on in vitro power production were also collected for largemouth bass (Micropterus salmoides) red muscle from previously published data on in vivo muscle activity. The in vivo data collected from swimming trout were similar to those for other species. The anterior red muscle of these fish has the longest duty cycle, the smallest phase shift between the onset of EMG activity and maximum muscle length during each tailbeat and undergoes the smallest strain or length change. For both trout and largemouth bass, work-loop experiments indicate that the majority of power for steady swimming is generated by the posterior muscle, as has been observed in other species.

Animals↗

Effects of sublethal ammonia exposure on swimming performance in rainbow trout (Oncorhynchus mykiss).

Adult trout Oncorhynchus mykiss fitted with a dorsal aortic catheter were exposed to 288+/-15 micromol l(-1) (mean +/- S.E.M.) total ammonia for 24h in water at a pH of 8.39+/-0.02, while swimming at a speed equivalent to 0.75 bodylengths s(-1) (BLs(-1)) in a Brett-type tunnel respirometer. The fish were then exposed to stepwise increments in swimming speed (0.25 BLs(-1) every 30 min) until exhaustion. Measurements of oxygen uptake (M(O2)) and plasma total ammonia levels and pH were made at each speed. Control trout were treated identically but without exposure to ammonia. Ammonia exposure caused an increase in plasma total ammonia level to 436+/-34 micromol l(-1), compared to 183+/-30 micromol l(-1)in control animals (N=6). A significant reduction in total plasma ammonia level was found in both groups during exercise, despite a large negative concentration gradient in those exposed to an elevated concentration of ammonia in water, which may indicate an active excretory process. The overall increase in plasma ammonia levels in exposed trout was associated with a significant reduction in critical swimming speed (U(crit)) to 1.61+/-0.17BL s(-1) from 2.23+/-0.15BL s(-1) in control animals. Ammonia-exposed trout had a significantly higher maintenance metabolic rate (MMR) than control fish, when estimated as the y-intercept of the relationship between swimming speed and M(O2). Active metabolic rate (AMR, maximum M(O2) as measured at U(crit)) was significantly lower in ammonia-exposed animals, leading to a profound reduction in factorial aerobic scope (AMR/MMR). Reduced U(crit) was also linked to a reduction in maximum tailbeat frequency. Calculation of membrane potentials (E(M)) in the white muscle of fish swum to U(crit) revealed a significant partial depolarisation of white muscle in ammonia-exposed fish. This may have prevented white muscle recruitment and contributed to the reduced maximum tailbeat frequency and overall impairment of swimming performance in the ammonia-exposed fish.

Ammonia↗

Swimming in needlefish (Belonidae): anguilliform locomotion with fins.

The Atlantic needlefish (Strongylura marina) is a unique anguilliform swimmer in that it possesses prominent fins, lives in coastal surface-waters, and can propel itself across the surface of the water to escape predators. In a laboratory flow tank, steadily swimming needlefish perform a speed-dependent suite of behaviors while maintaining at least a half wavelength of undulation on the body at all times. To investigate the effects of discrete fins on anguilliform swimming, I used high-speed video to record body and fin kinematics at swimming speeds ranging from 0.25 to 2.0 L s(-1) (where L is the total body length). Analysis of axial kinematics indicates that needlefish are less efficient anguilliform swimmers than eels, indicated by their lower slip values. Body amplitudes increase with swimming speed, but unlike most fishes, tail-beat amplitude increases linearly and does not plateau at maximal swimming speeds. At 2.0 L s(-1), the propulsive wave shortens and decelerates as it travels posteriorly, owing to the prominence of the median fins in the caudal region of the body. Analyses of fin kinematics show that at 1.0 L s(-1) the dorsal and anal fins are slightly less than 180 degrees out of phase with the body and approximately 225 degrees out of phase with the caudal fin. Needlefish exhibit two gait transitions using their pectoral fins. At 0.25 L s(-1), the pectoral fins oscillate but do not produce thrust, at 1.0 L s(-1) they are held abducted from the body, forming a positive dihedral that may reduce rolling moments, and above 2.0 L s(-1) they remain completely adducted.

Animals↗