Effects of linear acceleration on fish behavior and eye movements.

Behavioral responses and eye movements of fish during linear acceleration were reviewed. It is known that displacement of otoliths in the inner ear leads to body movements and/or eye movements. On the ground, the utriculus of the vestibular system is stimulated by otolith displacement caused by gravitational and inertial forces during horizontal acceleration of whole body. When the acceleration is imposed on the fish's longitudinal axis, the fish showed nose-down and nose-up posture for tailward and noseward displacement of otolith respectively. These responses were understood that the fish aligned his longitudinal body axis in a plane perpendicular to the direction of resultant force vector acting on the otoliths. When the acceleration was sideward, the fish rolled around his longitudinal body axis so that his back was tilted against the direction in which the inertial force acted on the otoliths. Linear acceleration applied to fish's longitudinal body axis evoked torsional eye movement. Direction of torsion coincided with the direction of acceleration, which compensate the change of resultant force vector produced by linear acceleration and gravity. Torsional movement of left and right eye coordinated with each other. In normal fish, both sinusoidal and rectangular acceleration of 0.1G could evoke clear eye torsion. Though the amplitude of response increased with increasing magnitude of acceleration up to 0.5 G, the torsion angle did not fully compensate the angle calculated from gravity and linear acceleration. Removal of the otolith on one side reduced the response amplitude of both eyes. The torsion angle evoked by rectangular acceleration was smaller than that evoked by sinusoidal acceleration in both normal and unilaterally labyrinthectomized fish. These results suggest that eye torsion of fish include both static and dynamic components.     

Computational methods for the analysis of swimming biomechanics

A series of computer programs is presented which enables the analysis of fish body shape and mass distributions, spine positions, spine curvatures and coordinates for the centre of mass. Data are derived from silhouette outlines of swimming fish, white muscle strains during swimming, white muscle force-time development functions for body bending cycles, muscle force and power production along the whole fish body and hydrodynamic efficiencies for fast-start swimming behaviours.     

Locomotion of Gymnarchus Niloticus： Experiment and Kinematics

In addition to forward undulatory swimming, Gymnarchus niloticus can swim via undulations of the dorsal fin while the body axis remains straight; furthermore, it swims forward and backward in a similar way, which indicates that the undulation of the dorsal fin can simultaneously provide bidirectional propulsive and maneuvering forces with the help of the tail fin. A high-resolution Charge-Coupled Device （CCD） imaging camera system is used to record kinematics of steady swimming as well as maneuvering in G. niloticus. Based on experimental data, this paper discusses the kinematics （cruising speed, wave speed, cycle frequency, amplitude, lateral displacement） of forward as well as backward swimming and maneuvering. During forward swimming, the propulsive force is generated mainly by undulations of the dorsal fin while the body axis remains straight. The kinematic parameters （wave speed, wavelength, cycle frequency, amplitude） have statistically significant correlations with cruising speed. In addition, the yaw at the head is minimal during steady swimming. From experimental data, the maximal lateral displacement of head is not more than 1% of the body length, while the maximal lateral displacement of the whole body is not more than 5% of the body length. Another important feature is that G. niloticus swims backwards using an undulatory mechanism that resembles the forward undulatory swimming mechanism. In backward swimming, the increase of lateral displacement of the head is comparatively significant; the amplitude profiles of the propulsive wave along the dorsal fin are significantly different from those in forward swimming. When G. niloticus does fast maneuvering, its body is first bent into either a C shape or an S shape, then it is rapidly unwound in a travelling wave fashion. It rarely maneuvers without the help of the tail fin and body bending.     

A continuous dynamic beam model for swimming fish

When a fish swims in water, muscle contraction, controlled by the nervous system, interacts with the body tissues and the surrounding fluid to yield the observed movement pattern of the body. A continuous dynamic beam model describing the bending moment balance on the body for such an interaction during swimming has been established. In the model a linear visco-elastic assumption is made for the passive behaviour of internal tissues, skin and backbone, and the unsteady fluid force acting on the swimming body is calculated by the 3D waving plate theory. The body bending moment distribution due to the various components, in isolation and acting together, is analysed. The analysis is based on the saithe (Pollachius virens), a carangiform swimmer. The fluid reaction needs a bending moment of increasing amplitude towards the tail and near-standing wave behaviour on the rear-half of the body. The inertial movement of the fish results from a wave of bending moment with increasing amplitude along the body and a higher propagation speed than that of body bending. In particular, the fluid reaction, mainly designed for propulsion, can provide a considerable force to balance the local momentum change of the body and thereby reduce the power required from the muscle. The wave of passive visco-elastic bending moment, with an amplitude distribution peaking a little before the mid-point of the fish, travels with a speed close to that of body bending. The calculated muscle bending moment from the whole dynamic system has a wave speed almost the same as that observed for EMG-onset and a starting instant close to that of muscle activation, suggesting a consistent matching between the muscle activation pattern and the dynamic response of the system in steady swimming. A faster wave of muscle activation, with a variable phase relation between the strain and activation cycle, appears to be designed to fit the fluid reaction and, to a lesser extent, the body inertia, and is limited by the passive internal tissues. Higher active stress is required from caudal muscle, as predicted from experimental studies on fish muscle. In general, the active force development by muscle does not coincide with the propulsive force generation on the tail. The stiffer backbone may play a role in transmitting force and deformation to maintain and adjust the movement of the body and tail in water.     

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.     

Effects of nutritional status on metabolic rate, exercise and recovery in a freshwater fish

The influence of feeding on swimming performance and exercise recovery in fish is poorly understood. Examining swimming behavior and physiological status following periods of feeding and fasting is important because wild fish often face periods of starvation. In the current study, researchers force fed and fasted groups of largemouth bass (Micropterus salmoides) of similar sizes for a period of 16 days. Following this feeding and fasting period, fish were exercised for 60 s and monitored for swimming performance and physiological recovery. Resting metabolic rates were also determined. Fasted fish lost an average of 16 g (nearly 12%) of body mass, while force fed fish maintained body mass. Force fed fish swam 28% further and required nearly 14 s longer to tire during exercise. However, only some physiological conditions differed between feeding groups. Resting muscle glycogen concentrations was twofold greater in force fed fish, at rest and throughout recovery, although it decreased in both feeding treatments following exercise. Liver mass was nearly three times greater in force fed fish, and fasted fish had an average of 65% more cortisol throughout recovery. Similar recovery rates of most physiological responses were observed despite force fed fish having a metabolic rate 75% greater than fasted fish. Results are discussed as they relate to largemouth bass starvation in wild systems and how these physiological differences might be important in an evolutionary context.     

On the mechanism of speed and altitude control in Drosophila melanogaster

ABSTRACT The total power output of tethered flying Drosophila melanogaster in still air depends on translational velocity components of image flow on the eye, whereas the orientation of the average flight force in the midsagittal plane of the fly is widely independent of visual input (Götz, 1968). The fly does not seem to control the vertical and the horizontal force component independently. Freely flying flies nevertheless generate different ratios between lift and thrust, simply by changing the inclination of their body. By the combined adjustment of the body angle and the total power output a fly appears to be able to stabilize height and speed (David, 1985). Here a possible mechanism is proposed by which the appropriate torque about the transverse body axis could be generated. Translational pattern motion influences the posture of the abdomen and the plane of wing oscillation. Thus the position of the centre of gravity relative to the flight force vector is changed. When abdomen and stroke plane deviate from an equilibrium state, a lever is generated by which the force vector will rotate the fly about its transverse axis.     

Liver-derived endocrine IGF-I is not critical for activation of skeletal muscle protein synthesis following oral feeding

Background

Like humans, fish can be classified according to their athletic performance. Sustained exercise training of fish can improve growth and physical capacity, and recent results have documented improved disease resistance in exercised Atlantic salmon. In this study we investigated the effects of inherent swimming performance and exercise training on disease resistance in Atlantic salmon. Atlantic salmon were first classified as either poor or good according to their swimming performance in a screening test and then exercise trained for 10 weeks using one of two constant-velocity or two interval-velocity training regimes for comparison against control trained fish (low speed continuously). Disease resistance was assessed by a viral disease challenge test (infectious pancreatic necrosis) and gene expression analyses of the host response in selected organs.

Results

An inherently good swimming performance was associated with improved disease resistance, as good swimmers showed significantly better survival compared to poor swimmers in the viral challenge test. Differences in mortalities between poor and good swimmers were correlated with cardiac mRNA expression of virus responsive genes reflecting the infection status. Although not significant, fish trained at constant-velocity showed a trend towards higher survival than fish trained at either short or long intervals. Finally, only constant training at high intensity had a significant positive effect on fish growth compared to control trained fish.

Conclusions

This is the first evidence suggesting that inherent swimming performance is associated with disease resistance in fish.     

Comparative biomechanical analysis of the rotational shot put technique

The study aimed to establish the modalities of the rotational shot put technique of two elite shot putters with substantially different constitutional characteristics. A biomechanical analysis of the technique was carried out using the APAS 3-D kinematic system, whereby a 15-segment model of shot putters was defined by 18 reference points. To enable the calculation of the kinematic and dynamic parameters, independent routines were programmed by the Matlab software. Anthropometric characteristics were established on the basis of 15 variables measured by the International Biological Programme (IBP) procedure. The results of the study revealed some differences between the athletes in terms of their mesomorphic constitutional component, body mass index, circular measures of the lower and upper extremities and the muscular, fat and bone mass. The technique models of both shot putters differ mostly in terms of the following kinematic and dynamic parameters: absolute release velocity, height of release, maximum angular velocity of the elbow of the throwing arm, trajectory of the centre of gravity of the body and the shot, torsional rotation of the shoulder axis relative to the hip axis, maximum force applied to the shot, kinetic energy and the kinetic energy differential of the shot.     

The contrasting effects of short-term climate change on the early recruitment of tree species

While interspecific competition is prevalent in natural systems, we do not yet understand how it can influence an individual’s phenotype within its lifetime and how this might affect performance. Morphology and swimming performance are two important fitness-related traits in fishes. Both traits are essential in acquiring and defending resources as well as avoiding predation. Here, we examined if interspecific competition could induce changes in morphology and affect the swimming performance of two strains of juvenile Atlantic salmon (Salmo salar). We imposed competitive scenarios on the fish using artificial streams containing different combinations of four interspecific competitors. Exposure to interspecific competitors induced morphological changes over time, through the development of deeper bodies, whereas controls free of interspecific competitors developed more fusiform body shapes. Furthermore, swimming performance was correlated to fusiform morphologies and was weaker for Atlantic salmon in competitive scenarios vs. controls. This implies that interspecific competition has direct effects on these fitness-related traits in Atlantic salmon. To the best of our knowledge, this is the first time that morphology, an important fitness-related trait linked to swimming performance, has been shown to be negatively impacted through interactions with an interspecific competitor.     

Reproductive ecology of cultured fish in the wild

Domestication has been shown to have an effect on morphology and behaviour of Atlantic salmon ( Salmo salar ). We compared swimming costs of three groups of juvenile Atlantic salmon subject to different levels of domestication: (1) wild fish; (2) first generation farmed fish origination from wild genitors; and (2) seventh generation farmed fish originating from Norwegian aquaculture stocks. We assessed swimming costs under two types of turbulent flow (one mean flow velocity of 23 cm s⁻¹ and two standard deviations of flow velocity of 5 and 8 cm s⁻¹). Respirometry experiments were conducted with fish in a mass range of 5–15 g wet at a water temperature of 15° C. Our results confirm (1) that net swimming costs are affected by different levels of turbulence such that, for a given mean flow velocity, fish spent 1·5‐times more energy as turbulence increased, (2) that domesticated fish differed in their morphology (having deeper bodies and smaller fins) and in their net swimming costs (being up to 30·3% higher than for wild fish) and (3) that swimming cost models developed for farmed fish may be also be applied to wild fish in turbulent environments.     

Differences in the energetic cost of swimming in turbulent flow between wild,farmed and domesticated juvenile Atlantic salmon Salmo salar

Domestication has been shown to have an effect on morphology and behaviour of Atlantic salmon (Salmo salar). We compared swimming costs of three groups of juvenile Atlantic salmon subject to different levels of domestication: (1) wild fish; (2) first generation farmed fish origination from wild genitors; and (2) seventh generation farmed fish originating from Norwegian aquaculture stocks. We assessed swimming costs under two types of turbulent flow (one mean flow velocity of 23 cm s^?1 and two standard deviations of flow velocity of 5 and 8 cm s^?1). Respirometry experiments were conducted with fish in a mass range of 5–15 g wet at a water temperature of 15° C. Our results confirm (1) that net swimming costs are affected by different levels of turbulence such that, for a given mean flow velocity, fish spent 1·5‐times more energy as turbulence increased, (2) that domesticated fish differed in their morphology (having deeper bodies and smaller fins) and in their net swimming costs (being up to 30·3% higher than for wild fish) and (3) that swimming cost models developed for farmed fish may be also be applied to wild fish in turbulent environments.     

A review of the multi-level adaptations for maximizing aerobic dive duration in marine mammals: from biochemistry to behavior

Marine mammals exhibit multi-level adaptations, from cellular biochemistry to behavior, that maximize aerobic dive duration. A dive response during aerobic dives enables the efficient use of blood and muscle oxygen stores, but it is exercise modulated to maximize the aerobic dive limit at different levels of exertion. Blood volume and concentrations of blood hemoglobin and muscle myoglobin are elevated and serve as a significant oxygen store that increases aerobic dive duration. However, myoglobin is not homogeneously distributed in the locomotory muscles and is highest in areas that produce greater force and consume more oxygen during aerobic swimming. Muscle fibers are primarily fast and slow twitch oxidative with elevated mitochondrial volume densities and enhanced oxidative enzyme activities that are highest in areas that produce more force generation. Most of the muscle mitochondria are interfibriller and homogeneously distributed. This reduces the diffusion distance between mitochondria and helps maintain aerobic metabolism under hypoxic conditions. Mitochondrial volume densities and oxidative enzyme activities are also elevated in certain organs such as liver, kidneys, and stomach. Hepatic and renal function along with digestion and assimilation continue during aerobic dives to maintain physiological homeostasis. Most ATP production comes from aerobic fat metabolism in carnivorous marine mammals. Glucose is derived mostly from gluconeogenesis and is conserved for tissues such as red blood cells and the central nervous system. Marine mammals minimize the energetic cost of swimming and diving through body streamlining, efficient, lift-based propulsive appendages, and cost-efficient modes of locomotion that reduce drag and take advantage of changes in buoyancy with depth. Most dives are within the animal’s aerobic dive limit, which maximizes time underwater and minimizes recovery time at the surface. The result of these adaptations is increased breath-hold duration and enhanced foraging ability that maximizes energy intake and minimizes energy output while making aerobic dives to depth. These adaptations are the long, evolutionary legacy of an aquatic lifestyle that directly affects the fitness of marine mammal species for different diving abilities and environments.     

Effects of satiation on feeding and swimming behaviour of planktivores

Hunger affects the feeding and swimming behaviour in fish. After 36 h of food deprivation, the feeding and swimming behaviour of Pseudorasbora parva (Cyprinidae) was studied under different prey densities (0.5, 1, 2, 5, 10 and 25 of Daphnia pulex per liter). The initial feeding rates showed marked variations in relation to prey availability. Under high prey densities, the initial feeding rate of fish was higher and subsequently decreased faster, when compared to those feeding under low prey densities. At higher prey densities, two factors were involved: that of higher prey encounter rates and also the attainment of food satiation at a faster rate. Across all prey densities, the feeding rates of fish reached a plateau after satiation. The swimming speed of fish was found to be negatively related to the prey density and a significant change in swimming speed was noted as being directly related to the level of satiation. It was found that the increasing satiation level greatly influenced the handling time and reactive volume of predator, which finally caused reduced feeding rates.     

The effects of body mass and feeding on metabolic rate in small juvenile Atlantic cod

Apparent specific dynamic action (SDA) amplitude in young juvenile Atlantic cod Gadus morhua (1 to 8 g wet mass), fed a standardized meal and then exercised in a circular swimming respirometer at a constant swimming speed of 0·5 ± 0·3 body lengths s^-1, occurred within l h after feeding in all juveniles. SDA amplitude were 1·4 to 1·8 times higher in fed fish compared to unfed fish, and rates of oxygen consumption decreased as body mass increased. SDA duration had a tendency to decrease with increasing body mass and was shortest, at 6 h, in the smallest fish (1–1·5 g), but increased to 10–11 h in the largest fish. Apparent SDA in fed fish ( R _r) scaled with a mass exponent of 0·89, while maximum metabolic rate ( R _max) determined by chasing fish to exhaustion and then measuring oxygen consumption for 12 h, and unfed routine metabolic rate (R_r) scaled with a mass exponent of 0·79 and 0·76 respectively. Relative aerobic scope ( R _max– unfed R _r) did not appear to vary over the 1 to 8 g increase in wet mass. These results show that as body mass increased in young juvenile Atlantic cod: (1) apparent SDA ( R _f) increased more rapidly than R _max, and (2) apparent SDA took up >98% of the relative aerobic scope and that young Atlantic cod allocated most of the energy to growth, and left little for other metabolic activities.     

Swimming restricted foraging behavior of two zooplanktivorous fishes <Emphasis Type="Italic">Pseudorasbora parva</Emphasis> and <Emphasis Type="Italic">Rasbora daniconius</Emphasis> (Cyprinidae) in a simulated structured environment

In littoral zones of aquatic systems, submerged macrophytes have marked structural variation that can modify the foraging activity of planktivores. Swimming and feeding behavior of Pseudorasbora parva and Rasbora daniconius (Cyprinidae) on their prey Daphnia pulex and Artemia salina, respectively, was studied in a series of laboratory experiments with varying stem densities. A range of stem densities was tested for each of the two species to compare the effect of simulated macrophytes on prey attack rates and swimming speed, average stem distance (D) was measured in fish body lengths for each of the two fish species. We found that, with reducing average stem distance, the attack rate decreased in the similar trend and this trend was similar for both fish species. However, the species differed in the degree to which swimming activity was hindered at increased stem densities, and this was due to species-specific differences in the distance moved with one tail beat. Therefore, we conclude that the reductions in swimming speed with reduced average stem distance are due to the differences in fish movement per tail beat.     

Migration speeds and orientation of Atlantic salmon and sea trout post-smolts in a Norwegian fjord system

We recorded the observed and actual swimming speeds of Atlantic salmon and sea trout post-smolts in a Norwegian fjord system, and initiated studies on the orientation mechanisms of the post-smolts. We tracked Atlantic salmon and sea trout with acoustic transmitters for up to 14 h after release. The actual swimming speed and direction of a fish relative to the ground is the vector sum of the observed movements of the fish and the movements of the water. We determined actual swimming speeds and directions of the post-smolts, which reflect their real swimming capacities and orientation, by corrections for the speed and direction of the water current. The post-smolts were actively swimming. The observed direction of movement was dependent on the actual movement of the fish and not the water current. Water currents were not systematically used as an orientation cue either in Atlantic salmon or sea trout, as the actual movements were random compared to the direction of the water current. The actual movement of sea trout were in all compass directions, with no systematic pattern. The Atlantic salmon also moved in all compass directions, but with the lowest frequency of actual movement towards the fjord.     

Locomotion of bluefish.

1. Pressure previously measured on the body surface of swimming bluefish were resolved into their backward vectorial components to allow calculation of profile drag. It was 0.18 kg at a speed of 1.8 m/sec. Tangential drag was calculated as if for a thin plate of an area equal to that of the fish. It was 0.08 kg at 1.8 m/sec. Net drag, 0.26 kg, was the sum of profile and tangential drag. 2. Thrust and drag also were calculated from the changes of acceleration measured during steady swimming, assuming that thrust took place only during the acceleration phase, whereas drag occurred during both acceleration and deceleration. This drag was 0.08 kg at a speed of 1.1 m/sec. It is compatible with the drag of 0.26 at 1.8 m/sec calculated from profile and tangential drag provided drag varies as the square of velocity. 3. The force required to produced maximal acceleration was measured during a scare. It was calculated to be 6.9 kg at a peak acceleration of 3 g. 4. The compression strength of th vertebrae was found to be approximately 20 kg per cm2, or roughly three times the force encountered during maximal acceleration. This safety factor of 3 would be reduced when the back was curved, or if opposing groups of muscles were under tension. 5. The finding that a bluefish can accelerate at 3 g and that the vertebral column is strongg enough to withstand this force indicates that the muscles and body structure of a bluefish would be able to withstand the force of gravity if the fish were otherwise equipped for terrestrial life. This fish may have evolved these strengths simultaneously with land animals. It is speculated that other fish may have evolved some degree of strength to overcome inertia and drag during aquatic locomotion, and this evolution may have been a prelude to terrestrial locomotion.     

Dietary fatty acid composition affects the repeat swimming performance of Atlantic salmon in seawater

Repeated critical swimming performance trials (Ucrit) were performed on Atlantic salmon (Salmo salar) to test the null hypothesis that the source of dietary lipids (fish-based, poultry-based, and plant-based) does not influence exercise and recovery performance. Four diets were prepared by extensively replacing supplemental lipid from anchovy oil (AO; 100% AO at 150 g/kg) with cold pressed flaxseed oil (FO; 25% AO, 75% FO), sunflower oil (SO; 25% AO, 75% SO), or poultry fat (PF; 25% AO, 75% PF). These diets had equivalent protein and energy concentrations, but due to the different supplemental lipid sources, varied widely in their fatty acid composition. Fish fed AO had a significantly higher (P<0.05) first Ucrit (2.62+/-0.07 body lenght s(-1)) than those fed PF (2.22+/-0.12 body lenght s(-1)) that had low muscle ratios of n-3 highly unsaturated fatty acids (n-3 HUFA) to saturated fatty acids (SFA) and arachidonic acid (AA), and high levels of oleic acid. Fish in the FO and SO diet groups swam as well as AO-fed fish in both swimming trials. The performance of fish fed AO decreased significantly (P<0.05) during the second swimming trial (i.e. Ucrit2/Ucrit1=0.92+/-0.02). No significant differences occurred between diet groups for the second swim trial. There was a positive correlation between both n-3 HUFA/SFA and n-3 HUFA/AA ratios, and Ucrit1. A negative correlation was found between dietary AA and oleic acids, and Ucrit1. The present study suggests that low dietary n-3 HUFA/ SFA and n-3 HUFA/AA ratios may negatively affect swimming performance. The former possibly can be offset by increasing linoleic acid in the presence of nutritionally adequate n-3 HUFA (e.g. SO diet). Lipid supplements consisting largely of vegetable oils did not compromise fish cardiorespiratory physiology under the conditions of this study.     

Acid–base and respiratory effects of confinement in Atlantic salmon affected with amoebic gill disease

Amoebic gill disease (AGD) in cultured salmonids causes severe multifocal hyperplastic lesions in the gills with the potential to influence respiratory and acid–base physiology. Atlantic salmon Salmo salar affected with AGD were surgically implanted with dorsal aortic catheters and, following recovery, were confined for 5 min ( n  = 16) or left undisturbed ( n  = 8). Confinement caused an acute extracellular acidosis that was corrected in 6 h amongst surviving fish. There was a gradual increase in plasma lactate concentrations that peaked at 1 h post-confinement then declined by 9 h recovery. In a second experiment, AGD-affected fish were confined then recovered either in a tank of static water ( n  = 9) or while being forced to swim at 1·5 body lengths s⁻¹ ( n  = 6). There was no significant difference between fish recovered by swimming and those in static water in terms of recovery of the acute extracellular acidosis and lactate accumulations coincident with exhaustive exercise. Confinement severely compromised the survival of AGD-affected Atlantic salmon, although survivors appeared to recover similarly to other studies. Forced swimming of AGD-affected Atlantic salmon following confinement did not facilitate recovery and is unlikely to be a useful strategy for mitigating the effects of stressful episodes such as crowding and fish movement and commercial handling.