The effect of fasting and refeeding on temperature preference,activity and growth of roach, <Emphasis Type="Italic">Rutilus rutilus</Emphasis>

Most fish species are regularly subjected to periods of starvation during which a reduction of energy turnover might be favourable for the animal. This reduction of energy flux may be achieved by changes in thermal behaviour and/or swimming activity. We investigated such behavioural changes during starvation and subsequent refeeding in roach, Rutilus rutilus, with respect to energetic benefits and growth maximisation. Roach, acclimated to a wide range of temperatures (4, 12, 20, 24, 27 and 30 °C), were fed to excess, subjected to 3 weeks of starvation and subsequently refed in order to determine the temperature dependence of feeding rates, growth rates and conversion efficiency (K₁) under control conditions and during compensatory growth. When exposed to a thermal gradient, control animals preferentially selected a temperature of 26.8ǂ.9 °C, which is in the range of the optimal temperatures for feeding, growth and conversion efficiency. Starving fish showed a distinct circadian pattern of the mean selected temperature (MST). They migrated to cooler water in the dark (MST_dark=22.8ǃ.1 °C) but returned to warmer water during daytime. This behaviour may be regarded as a trade-off between the potentially higher food density in warmer water areas and the energetic benefit of selecting cooler water patches. The circadian pattern of MST was gradually abandoned upon refeeding and control values were reached again after 3 weeks. Energetically more effective than behavioural hypothermia was the reduction of swimming activity. During starvation, activity peaks were slightly lower than under control conditions and mean daily activity decreased by about 50%. Swimming velocity, however, was not affected by feeding regime. After a period of starvation fish showed compensatory growth at all temperatures, even below 12 °C, where these animals normally do not grow. This suggests that after a period of starvation the critical temperature for growth shifts to lower values.     

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

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

Use of a self-propelled,fish-shaped,computer model to study the effects of shape and tail-beat frequency on torque and velocity in salmon-shaped fish

The relationship between fish shape, swimming ability and energy consumption during swimming in fish is complex and not well understood. In this paper, we show how a self-propelled 3-D fish model can be used to examine the effect of controlled changes in some shape parameters. Parameters of the model fish are modified and the resulting fish activated for short swimming episodes during which swimming velocity, torque and energy expenditure are calculated in the computer environment. The effect of shape was determined for two different fish shapes swimming at three different tail-beat frequencies (1.43, 0.94 and 0.64?Hz). The simulation results indicate that fish model one (based on a salmon) has stronger swimming ability than fish model two (a modified salmon fish shape) even though energy expenditure of fish shape two is greater than that of fish shape one. In the same fish types, the fish-swimming velocity and energy expenditure are proportional to tail-beat frequency. This model has the potential to be useful, particularly for predicting fish behavior in fish swim ways and the tail-water of energy turbines.     

The size and conjugation of change of parameters of growth and energetics of fish caused by oscillation of temperature

Small deviations of temperature from stationary optimum values optimize the growth and energetics of fish. The positive influence on the growth and energetics of fish at fluctuations of temperature is more expressed when they occur as a result of spontaneous swimming in the temperature gradient field. Changes in growth rates, respiration intensity, and consumption of energy per unit of weight that occur under the influence of oscillation of temperature are closely correlated with each other.     

Diurnal Variations in Swimming Activity of Rutilus rutilus (Cyprinidae) in a Group under Tank Conditions

Measurements of the swimming activity of a group of roach (12–19 cm TL, average) in a circular swimming chamber revealed two distinct activity patterns: a diurnal and a nocturnal one. The experiments showed that, having the choice, two factors stimulated the rhythmicity of the swimming behaviour of the fish, i.e. light intensity and the presence of a current field in the proximity of the fish. During daytime (bright light conditions) the fish moved into the current field and swam on average at 0.4 BL/s (resting swimming). The roach remained swimming at this speed even if no current field was established, however, then distributed evenly in the basin. By contrast, during night (dim light conditions) the fish predominantly chose the still water section but swam on average with a cruising speed of 1.6 BL/s (night swimming). Accordingly, they did not seek the still water section for night swimming if the light was kept on. Then again, the fish distributed more or less evenly in the basin. The results support the hypothesis that the fish migrate during night-time and do this preferably in still water.     

Angling-induced cardiac disturbance of free-swimming largemouth bass (Micropterus salmoides) monitored with heart rate telemetry

The sub‐lethal effects of catch‐and‐release angling have been poorly studied because of the difficulties in monitoring physiological parameters in free‐swimming fish. Laboratory studies provide the opportunity to examine sub‐lethal effects in controlled environments, but do not incorporate site‐specific characteristics. In this study we angled free‐swimming largemouth bass (Micropterus salmoides) equipped with heart rate transmitters to exhaustion using rod and reel, and exposed fish to air for 30 s. Experiments were repeated at four water temperatures (13, 17, 21, and 25°C). These field data were compared with published findings from largemouth bass collected at the same water temperatures in a controlled laboratory setting using Doppler flow probes. Field collected heart rate data increased with increasing water temperatures (Q₁₀ values 1.30–1.37). Pre‐disturbance heart rates were ～30% higher for free‐swimming fish in the field than previously collected laboratory data at the same water temperatures. Fish angled in the field exhausted ～40% more rapidly than fish chased in the laboratory. Maximal heart rate was ～15% higher for free‐swimming fish in the field than for data collected from laboratory restrained fish, but scope for heart rate was reduced by up to 20% in the field, especially at higher water temperatures. Heart rate in free‐swimming fish was highly variable at all times, obscuring clear recovery patterns. Conversely, laboratory cardiac parameters exhibited less variable patterns, peaking clearly following disturbances and recovering in about 135 min, independent of water temperature. Based upon these findings, we suggest that comprehensive studies incorporating both laboratory and field experiments are needed for truly understanding the effect of catch‐and‐release angling on fish.     

Swimming Behavior Analysis Based on Bacterial Chemotaxis in Solution

Microrobots is playing more and more important roles for medical applications,such as targeting tumoral lesions for therapeutic purposes,Minimally Invasive Surgery (MIS) and highly localized drug delivery.However,energy efficient propulsion system poses significant challenges for the implementation of such mobile robots.Flagellated chemotactic bacteria can be used as an effective integrated propulsion system for microrobots.In this paper,we proposed a new type of propulsion method that is inspired by the motility mechanism of flagellated chemotactic bacteria in different pH gradients.The pH gradient field was established in solution through electrolysis method.The distribution of the pH values in solution was measured with pH indicator and analyzed with image processing technology,and the mechanism by which the pH values changed was also discussed.The swimming speed and direction of the bacteria were studied experimentally.Through analyzing the key parameters,such as stabilization time and electrode voltage,the optimal design of propulsion mechanism based on bacteria motion in the pH gradient field was proven.     

鱼类通过鱼道内水流速度障碍能力的评估方法

鱼类通过鱼道内水流速度障碍能力的量化对鱼道设计有重要理论和实际价值,其基础是鱼类游泳能力的测定.首先对鱼类游泳能力的研究方法进行了概述总结,指出了鱼类游泳能力经典测试方法存在测定流场与自然情况相差较大的不足;分析了关键要素如鱼类行为特征、生理耗能规律及水力特性对鱼类通过水流速度障碍能力的影响;提出了分析鱼类游泳行为和能力与特征流场的关系,探讨鱼类通过水流障碍行为规律和生理疲劳恢复特征,通过研究仿自然流态下的鱼类自由游泳行为、水力计算及生理耗能的关系,构建多因素鱼类游泳能力关系式,定量评价鱼类通过鱼道内水流速度障碍的发展方向.     

Assessing the ecological relevance of swimming performance traits: a case study of bluegill sunfish (<Emphasis Type="Italic">Lepomis macrochirus</Emphasis>)

A variety of fish species show habitat-related variation in traits associated with swimming performance and foraging behavior. This commonly manifests as a distinction between open water and shallow water littoral ecotypes. In bluegill sunfish (Lepomis macrochirus), open water fish exhibit greater energy economy and speed during sustained locomotion than those from the littoral, whereas littoral fish were more maneuverable than their open water counterparts. These distinctions are associated with variation in diet and foraging behavior and may represent a resource polyphenism that enhances fitness through more effective exploitation of particular habitat types. A lack of field data means that polyphenisms have not been placed in context with swimming behavior in the field. We have used 3D videography to quantify bluegill field swimming performance in open water and littoral habitats. This revealed patterns of performance variation that parallel the trait variation previously established in the laboratory. Open water fish utilized faster average swimming speeds than inshore fish, while indicators of nonlinearity and unsteadiness were greater in the littoral fish. There are, however, differences in propulsive behavior between the field and laboratory. Pectoral-fin-powered, median-paired fin swimming is rarely employed by open water fish. Field body-caudal fin swimming involves short sequences of propulsive tail beats interspersed with gliding, rather than the repeated propulsive cycles employed under steady-state conditions. This suggests a need to re-evaluate the applicability of steady-state performance traits to behavior and fitness in the field and highlights the general importance of obtaining field performance data.     

New insights into fish swimming: a proteomic and isotopic approach in gilthead sea bream

Moderate exercise enhances fish growth, although underlying physiological mechanisms are not fully known. Here we performed a proteomic and metabolic study in white (WM) and red (RM) muscle of gilthead sea bream juveniles swimming at 1.5 body lengths per second. Continuous swimming for four weeks enhanced fish growth without increasing food intake. Exercise affected muscle energy stores by decreasing lipid and glycogen contents in WM and RM, respectively. Protein synthesis capacity (RNA/protein), energy use (estimated by lipid-δ(13)C and glycogen-δ(13)C), and enzymatic aerobic capacity increased in WM, while protein turnover (expressed by δ(15)N-fractionation) did not change. RM showed no changes in any of these parameters. 2D-PAGE analysis showed that almost 15% of sarcoplasmic protein spots from WM and RM differed in response to exercise, most being over-expressed in WM and under-expressed in RM. Protein identification by MALDI-TOF/TOF-MS and LC-MS/MS revealed exercise-induced enhancement of several pathways in WM (carbohydrate catabolism, protein synthesis, muscle contraction, and detoxification) and under-expression of others in RM (energy production, muscle contraction, and homeostatic processes). The mechanism underpinning the phenotypic response to exercise sheds light on the adaptive processes of fish muscles, being the sustained-moderate swimming induced in gilthead sea bream achieved mainly by WM, thus reducing the work load of RM and improving swimming performance and food conversion efficiency.     

Shape variation in a benthic stream fish across flow regimes

Evolution of fish body shapes in flowing and non-flowing waters have been examined for several species. Flowing water can select for fish body shapes that increase steady swimming efficiency, whereas non-flowing water can favor shapes that increase unsteady swimming efficiency. Benthic stream fishes often use areas near the substrate that exhibit reduced or turbulent flow, thus it is unclear which swimming forms would be favored in such environments, and how shape might change across flow regimes. To test the relationship between fish body shape and flow regime in a benthic stream fish, we used geometric morphometric techniques to characterize lateral body shape in mountain sucker (Catostomus platyrhynchus) across flow rates, using stream gradient as an indicator of stream flow. Mountain suckers from low-flow environments were more streamlined, consistent with steady swimming body shapes, whereas mountain suckers from high flows had deeper bodies, consistent with unsteady swimming body shapes. In addition, smaller individuals tended to have more robust body shapes. These patterns are opposite to those predicted for stream fishes in the mid-water column. The benthic stream environment represents a distinct selective environment for fish shape that does not appear to conform to the simple dichotomy of flowing versus non-flowing water.     

Optimal swimming speed in head currents and effects on distance movement of winter-migrating fish

Migration is a commonly described phenomenon in nature that is often caused by spatial and temporal differences in habitat quality. However, as migration requires energy, the timing of migration may depend not only on differences in habitat quality, but also on temporal variation in migration costs. Such variation can, for instance, arise from changes in wind or current velocity for migrating birds and fish, respectively. Whereas behavioural responses of birds to such changing environmental conditions have been relatively well described, this is not the case for fish, although fish migrations are both ecologically and economically important. We here use passive and active telemetry to study how winter migrating roach regulate swimming speed and distance travelled per day in response to variations in head current velocity. Furthermore, we provide theoretical predictions on optimal swimming speeds in head currents and relate these to our empirical results. We show that fish migrate farther on days with low current velocity, but travel at a greater ground speed on days with high current velocity. The latter result agrees with our predictions on optimal swimming speed in head currents, but disagrees with previously reported predictions suggesting that fish ground speed should not change with head current velocity. We suggest that this difference is due to different assumptions on fish swimming energetics. We conclude that fish are able to adjust both swimming speed and timing of swimming activity during migration to changes in head current velocity in order to minimize energy use.     

Energy utilization during swimming and cost of locomotion in larval and juvenile fish

Oxygen consumption and ammonia excretion rates increased in an accelerated manner in larvae and juveniles of whitefish (Coregonus sp.) as a function of swimming speed. The three-dimensional patterns of fish metabolic rates (expressed as energy consumed or nitrogen excreted) versus body weight and swimming speed show that the total standard metabolic rate (i. e. at extrapolated zero swimming speed) increased during early life of whitefish, followed by the expected decrease. This phenomenon might be due to the profound changes in oxidative and glycolytic enzyme activities during fish “metamorphosis”. Standard metabolic rate of ammonia excretion, as a principal product of protein catabolism in fish, decreased by one order of magnitude in early coregonid ontogenesis. This means that protein utilization as an energy source decreases as far as standard metabolism is concerned, but increases with swimming speed. This trend is opposite that in adult fish, where protein utilization in the overall energy supply is diminished at increasing swimming speed. The cost of locomotion offish larvae and juveniles demonstrates that the energy expenditure increases logarithmically with decreasing fish size but at an accelerated rate as compared to adult fish. This contradicts earlier estimates of lower cost of swimming in fish larvae than cost of paddle-propulsion swimming in small invertebrates or cost of flying in insects.     

不同游泳速度条件下瓦氏黄颡幼鱼的有氧和无氧代谢反应

在(25±1)℃的条件下,测定瓦氏黄颡(Pelteobagrus vachelli Richardson)幼鱼体重(4.34±0.13)g的临界游泳速度(Ucrit),然后分别以临界游泳速度的不同百分比(20、40、60、80、100%Ucrit)将实验鱼分为5个速度处理组,另外设置静止对照组和高速力竭对照组。处理组实验鱼在不同游泳速度下分别游泳20min,在此过程中测定并计算运动代谢率(Activity metabolic rate,AMR),随后测定肌肉、血液和肝脏中的乳酸、糖原和葡萄糖含量。结果显示:实验鱼的绝对临界游泳速度为(48.28±1.02)cm/s,相对临界游泳速度为(6.78±0.16)BL/s;随着游泳速度的提高AMR显著增加(Pcrit时肌乳酸和血乳酸含量显著高于80%Ucrit的水平(P0.05);100%Ucrit时肝糖原含量显著低于40%Ucrit的水平(P0.05)。经计算瓦氏黄颡幼鱼到达临界游泳速度时的无氧代谢功率比例仅为11.0%,表明其游泳运动主要以有氧代谢供能;实验鱼的无氧代谢大约在80%Ucrit才开始启动,与其他鱼类比较启动时间较晚,说明其游泳运动对无氧代谢的依赖程度较低。研究提示瓦氏黄颡幼鱼是一种有氧运动能力较强的鱼类,这一能量代谢特征可能与提高其生存适合度有关。       

Swimming performance and energy homeostasis in juvenile laboratory raised fathead minnow (Pimephales promelas) exposed to uranium mill effluent

Research at the Key Lake uranium mill (Saskatchewan, Canada) suggests effluent discharged from the mill affects energy stores of resident fish, but the mechanisms by which energy homeostasis is affected and the subsequent effects on swimming performance are unknown. In the present study larvae were collected from laboratory raised adult fathead minnow (Pimephales promelas) exposed to 5% diluted uranium mill effluent or control (dechlorinated municipal) water, and reared in the same treatments to 60 days post hatch (dph). Critical swimming speed (U_crit) was significantly lower in effluent exposed 60 dph fish compared to control fish. Fish used in tests were considered fatigued and compared to fish without swim testing (non-fatigued). There were no differences in whole body glycogen or triglyceride concentrations between effluent exposed versus control fish. However, fatigued fish from both treatments had significantly lower triglycerides, but not glycogen, compared to non-fatigued fish from the same treatment. Whole body β–hydroxyacyl coenzymeA dehydrogenase activity was similar in fish from both treatments, but citrate synthase activity was significantly lower in effluent exposed fish. Our results suggest uranium mill effluent exposure in the laboratory affects aerobic energy metabolism and swimming performance in juvenile fathead minnow, which could affect wild fish survivability.     

Water temperature, light intensity and zooplankton density and the feeding activity of juvenile brook charr (Salvelinus fontinalis)

SUMMARY 1. The objective of this study was to evaluate the effects of zooplankton biomass (as a measure of density), fish biomass, light intensity and water temperature on the attack rate and swimming characteristics (i.e. swimming speed and angle of turn) of juvenile (1+) brook charr (Salvelinus fontinalis) in field enclosures. We used a portable underwater camera system in a series of pelagic enclosures to quantify the feeding behaviour of brook charr over a gradient of natural conditions. 2. In simple linear or non‐linear regression models we found (i) that attack rate and angle of turn were positively related to water temperature, (ii) that attack rate and swimming speed were positively related to zooplankton biomass and light intensity and (iii) that attack rate was positively related to swimming speed. In multiple regression models, fish biomass, light intensity and variance of the angle of turn accounted for 87% of the variation in attack rate. Light intensity and water temperature accounted for 86% of the variation in swimming speed. Fish gut fullness and attack rate accounted for 83% of the variation in the variance of the angle of turn executed by fish. 3. The increase in the number of attacks as zooplankton biomass increases conforms to the general positive functional response observed in other fish species. Our results also support the hypothesis that swimming speed increases with prey biomass. We did not observe a plateau in attack rate as zooplankton biomass increased. As our experiments were performed under natural biotic and abiotic conditions, factors other than zooplankton biomass might affect or limit this response, such as water temperature and light intensity. 4. Because zooplankton biomass was correlated with water temperature and light intensity, it was not possible to evaluate the independent contribution of these factors on the attack rate and swimming characteristics (swimming speeds and angle of turn) of brook charr. However, this study highlighted the impact of these factors on the feeding behaviour of juvenile brook charr when feeding in the pelagic habitat under natural conditions, and their importance in future models of optimal foraging and fish habitat quality.     

Automated Reconstruction of Three-Dimensional Fish Motion,Forces, and Torques

Fish can move freely through the water column and make complex three-dimensional motions to explore their environment, escape or feed. Nevertheless, the majority of swimming studies is currently limited to two-dimensional analyses. Accurate experimental quantification of changes in body shape, position and orientation (swimming kinematics) in three dimensions is therefore essential to advance biomechanical research of fish swimming. Here, we present a validated method that automatically tracks a swimming fish in three dimensions from multi-camera high-speed video. We use an optimisation procedure to fit a parameterised, morphology-based fish model to each set of video images. This results in a time sequence of position, orientation and body curvature. We post-process this data to derive additional kinematic parameters (e.g. velocities, accelerations) and propose an inverse-dynamics method to compute the resultant forces and torques during swimming. The presented method for quantifying 3D fish motion paves the way for future analyses of swimming biomechanics.     

Phenotypic plasticity and predator effects on morphology and physiology of crucian carp in nature and in the laboratory

Crucian carp Carassius carassius show great phenotypic plasticity in individual morphology and physiology, and strong variation in population density in different fish communities. Small fish with shallow bodies and large heads are typical in overcrowded monospecific fish communities in small ponds, whereas deep-bodied, large fish are found in larger, multispecies lakes. Crucian carp are especially vulnerable to predation by piscivorous fish and their greater relative body depth in multispecies fish communities has been proposed to be an induced defence against size-limited predation, and hence to be an adaptive feature. Data are presented here on the two divergent body forms in field populations in eastern Finland, together with results of laboratory experiments on predator effects on morphology and physiology (growth, respiration, heart rate). The deep body can be achieved in a few months by introducing a low population density of shallow-bodied fish into a food-rich environment with no piscivores. In the laboratory, both the presence of piscivores (chemical cues) and enhanced food availability increased the relative depth of crucian carp, but only to a modest extent when compared to natural variation. It is concluded that the deep-body form of crucian carp in the low density populations of multispecies fish communities is the normal condition. Reproduction in monospecific ponds results in high intraspecific competition, low growth rate and a stunted morphology. According to pilot tests, the mechanism behind the predator effect in the laboratory might be a behavioural reaction to chemical cues (alarm substances/predator odour) causing changes in energy allocation: predator-exposed crucian carp adopt a 'hiding' mode with decreased activity (less swimming, lower respiration and heart rate) and with higher overall growth. Whether, and to what extent, this predator-induced mechanism works in nature is unclear.     

Individual variations and interrelationships in feeding rate,growth rate,and spontaneous activity in hybrid tilapia (Oreochromis niloticus × O. aureus) at different feeding frequencies

A 30‐day growth trial was conducted to investigate the individual variations in feeding rate, growth rate, spontaneous activity, and their possible interrelationships in hybrid tilapia (Oreochromis niloticus × O. aureus) reared individually at feeding frequencies of one and two meals daily. Results show that the feeding rate in energy (FRe), specific growth rate in energy (SGRe), and food conversion efficiency (FCE) of fish fed twice a day were significantly higher than that of fish fed only once. However, no significant differences in distance or in time spent swimming were observed between the groups. SGRe was positively correlated with FRe in fish fed only once daily, and SGRe was positively correlated with food conversion efficiency in energy (FCEe) between the two groups. SGRe was only inversely correlated with the distance that fish swam when fed once daily. The results suggest that the individual growth differences of hybrid tilapia could be attributed mainly to variations in FRe, and that the energy costs of spontaneous activity play an important role in the differences at feeding frequency of one meal a day. However, at feeding frequencies of two meals a day, individual growth differences in fish may be caused by variations in FCEe, and energy costs of spontaneous activity have only marginal effects on the differences.     

Physiological Plasticity to Water Flow Habitat in the Damselfish,Acanthochromis polyacanthus: Linking Phenotype to Performance

The relationships among animal form, function and performance are complex, and vary across environments. Therefore, it can be difficult to identify morphological and/or physiological traits responsible for enhancing performance in a given habitat. In fishes, differences in swimming performance across water flow gradients are related to morphological variation among and within species. However, physiological traits related to performance have been less well studied. We experimentally reared juvenile damselfish, Acanthochromis polyacanthus, under different water flow regimes to test 1) whether aspects of swimming physiology and morphology show plastic responses to water flow, 2) whether trait divergence correlates with swimming performance and 3) whether flow environment relates to performance differences observed in wild fish. We found that maximum metabolic rate, aerobic scope and blood haematocrit were higher in wave-reared fish compared to fish reared in low water flow. However, pectoral fin shape, which tends to correlate with sustained swimming performance, did not differ between rearing treatments or collection sites. Maximum metabolic rate was the best overall predictor of individual swimming performance; fin shape and fish total length were 3.3 and 3.7 times less likely than maximum metabolic rate to explain differences in critical swimming speed. Performance differences induced in fish reared in different flow environments were less pronounced than in wild fish but similar in direction. Our results suggest that exposure to water motion induces plastic physiological changes which enhance swimming performance in A. polyacanthus. Thus, functional relationships between fish morphology and performance across flow habitats should also consider differences in physiology.