Effect of repeated exercise fatigue on the swimming performance of juvenile rock carp (Procypris rabaudi,Tchang)

The swimming performance of juvenile rock carp (Procypris rabaudi, Tchang) subjected to repeated fatigue exercise was studied using a flume-type respirometer at 20°C. The critical swimming speed (U_crit) and oxygen consumption rate (MO₂) of juvenile rock carp were measured during two successive stepped velocity tests, following a 60 min rest interval. U_crit of rock carp was giving a recovery ratio (R_r) of 92.64%, and exertion exercise decreases U_crit. When MO₂ was plotted as a linear function of U, the slope for trial 1 was 1.06 and 1.50 for trial 2, indicating a decreasing in swimming efficiency. The maximum metabolic rate (MMR) increased from 17.06 ± 1.14 mmol O₂/(kg·hr) to 19.14 ± 1.23 mmol O₂/(kg·hr), and the exercise post oxygen consumption rate (EPOC) increased from 9.00 to 9.65 mmol O₂/kg. Repeated fatiguing exercise increased both the aerobic and anaerobic cost of reaching U_crit, but anaerobic metabolism accounted for a larger proportion in the trial 2. The data investigation on the swimming performance and the physiological response to fatigue provide important design criteria for fishways.     

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.     

Hypoxia tolerance variance between swimming and resting striped bass Morone saxatilis

Individual striped bass Morone saxatilis were each exposed in random order to aquatic hypoxia (10% air saturation) either while swimming at 50% of their estimated critical swimming speed (U_crit) or while at rest until they lost equilibrium. Individuals were always less tolerant of hypoxia when swimming (P < 0·01); the average fish was over five times more tolerant to the same hypoxia exposure when not swimming. There was no relationship between an individual's rank order of hypoxia tolerance (HT) under the two flow regimes, suggesting that different factors determine an individual's HT when at rest than when swimming.     

An interspecific comparison between morphology and swimming performance in cyprinids

Flow regimes are believed to be of major evolutionary significance in fish. The flow regimes inhabited by cyprinids vary extensively from still flow regimes to riptide flow regimes. To test (i) whether flow‐driven swimming performance and relevant morphological differentiation are present among fish species and (ii) whether evolutionary shifts between high‐flow and low‐flow habitats in cyprinids are associated with evolutionary trade‐offs in locomotor performance, we obtained data on both steady and unsteady swimming performance and external body shape for 19 species of cyprinids that typically occur in different flow regimes (still, intermediate and riptide). We also measured the routine energy expenditure (RMR) and maximum metabolic rate (MMR) and calculated the optimal swimming speed. Our results showed that fish species from riptide groups tend to have a higher critical swimming speed (U_crit), maximum linear velocity (V_max) and fineness ratio (FR) than fish from the other two groups. However, there was no correlation between the reconstructed changes in the steady and unsteady swimming performance of the 19 species. According to the phylogenetically independent contrast (PIC) method, the U_crit was actively correlated with the MMR. These results indicated that selection will favour both higher steady and unsteady swimming performance and a more streamlined body shape in environments with high water velocities. The results suggested that steady swimming performance was more sensitive to the flow regime and that for this reason, changes in body shape resulted more from selective pressure on steady swimming performance than on unsteady swimming performance. No evolutionary trade‐off was observed between steady and unsteady swimming performance, although U_crit and MMR were found to have coevolved. However, a further analysis within each typically occurring habitat group suggested that the trade‐off that may exist between steady and unsteady swimming performance may be concealed by the effect of habitat.     

Ontogenetic differentiation of swimming performance in Gilthead seabream (Sparus aurata, Linnaeus 1758) during metamorphosis

The critical swimming speed (U_crit) of gilthead seabream (Sparus aurata, Linnaeus 1875) was studied in two ontogenetic phases, early (13.7-18.7 mm total length, TL) and late metamorphosis (20.4-34.3 mm TL, after the full development of fin meristics and during squamation ontogeny), under four exercise temperatures (15, 20, 25 and 28 °C). Both the exercise temperature and the ontogenetic stage had a significant effect on the relative U_crit (RU_crit) of S. aurata, with the fish of early metamorphosis phase (E group) presenting significantly higher RU_crit than those of the late metamorphosis stage (L group). This ontogenetic shift in swimming performance was accompanied by significant ontogenetic shifts of body shape and of muscle anatomy. Compared to the L group, S. aurata of the E group were characterized by a streamline body shape and significantly higher relative contribution of the slow-red muscle to the cross-sectional area of the body (31.0 ± 1.3% vs 12.0 ± 1.2% in the L group).     

Size-related effects and the influence of metabolic traits and morphology on swimming performance in fish

Energy metabolism fuels swimming and other biological processes. We compared the swimming performance and energy metabolism within and across eight freshwater fish species. Using swim tunnel respirometers, we measured the standard metabolic rate (SMR) and maximum metabolic rate (MMR) and calculated the critical swimming speed (U_crit). We accounted for body size, metabolic traits, and some morphometric ratios in an effort to understand the extent and underlying causes of variation. Body mass was largely the best predictor of swimming capacity and metabolic traits within species. Moreover, we found that predictive models using total length or SMR, in addition to body mass, significantly increased the explained variation of U_crit and MMR in certain fish species. These predictive models also underlined that, once body mass has been accounted for, U_crit can be independently affected by total length or MMR. This study exemplifies the utility of multiple regression models to assess within-species variability. At interspecific level, our results showed that variation in U_crit can partly be explained by the variation in the interrelated traits of MMR, fineness, and muscle ratios. Among the species studied, bleak Alburnus alburnus performed best in terms of swimming performance and efficiency. By contrast, pumpkinseed Lepomis gibbosus showed very poor swimming performance, but attained lower mass-specific cost of transport (MCOT) than some rheophilic species, possibly reflecting a cost reduction strategy to compensate for hydrodynamic disadvantages. In conclusion, this study provides insight into the key factors influencing the swimming performance of fish at both intra- and interspecific levels.     

Effects of lateral morphology on swimming performance in two sturgeon species

Fish express a high degree of diversity in morphology, which is closely related to behaviors such as swimming ability. The effect of morphology on swimming performance is explored using geometric morphometric analyses and classic critical swimming speed (U_crit) tests in Chinese sturgeon Acipenser sinensis and Siberian sturgeon A. baerii. It was found that A. sinensis is a stronger swimmer compared to A. baerii, with an average 25% higher U_crit (expressed in body lengths per second). In A. sinensis, the depth and length of the snout and the trailing edge length of the dorsal fin were negatively correlated with U_crit, whereas the height of the trunk anterior, the leading edge length of the dorsal fin and anal fin, and the length and width of the ventral lobe were positively related to U_crit; similar relationships between U_crit and morphological characters of the anterior trunk, dorsal fin, anal fin and caudal fin were found in A. baerii. Moreover, although the degree of upward bending of the snout of A. baerii was negatively related to U_crit, there was a positive relationship between the length of the caudal peduncle and U_crit as well as between the dorsal tail lobe and U_crit. In addition, the streamline index (SI) was calculated by comparing landmark coordinates on the trunk displayed in the relative warp, with its corresponding point on the NACA (the U.S. National Advisory Committee for Aeronautics) airfoil shape. SI showed that the body shape in RW1 of the A. baerii with more swimming capacity was more approximate to the NACA 0016 airfoil shape, but there was no such symmetry for A. sinensis, possibly due to body bending caused by stiffness.     

Swimming performance of 12 Schizothoracinae species from five rivers

A series of stepped velocity tests were carried out in a Brett‐type swimming respirometer and the overall range in swimming performance for 12 Schizothoracinae species was measured. The relative critical swimming speed U_crit and burst speed U_burst decreased with body length, while absolute U_crit and U_burst increased with body length. U_crit increased with temperature up to approximately 15^° C and then decreased. Species with a high U_crit also displayed a higher U_burst.     

High levels of metacercarial infestation (family: Diplostomidae) do not affect host energetics and swimming performance in the Epaulette goby (Coryogalops sordidus,Gobiidae)

Parasites have deleterious effects on their hosts, often resulting in altered host behavior or increased energy expenditure. When organisms are exposed to suboptimal environments, parasite loading may increase. Microbialite pools along the warm temperate South African coastline have been hypothesized as refugia for Epaulette gobies (Coryogalops sordidus, Gobiidae) when they are outside of their previously known subtropical distribution. The aim of this study was to determine if C. sordidus individuals infected with metacercarial cysts display higher metabolic rates or different swimming behavior compared to noninfected individuals. We measured each goby's swimming performance using a critical station-holding speed (U_crit) test (n = 60) and visually scored their swimming behavior (n = 52) during these measurements. Also, we measured the metabolic rate of gobies using an intermittent flow respirometer system to determine standard metabolic rate (SMR) and maximum metabolic rate (MMR) from gobies at 21°C before and after swimming trials. Metacercarial load carried by infected gobies seemingly had no impact on the host's energetics (SMR or MMR), swimming ability (as repeated U_crit tests), or swimming behavior compared to noninfected gobies. Thus, the metacercarial intensity observed in gobies in the current study appeared to have no impact on host swimming performance or behavior. Furthermore, the swimming capacity observed for C. sordidus, in general, suggests that this goby is a poor swimmer compared to other gobiid species.     

Swimming performance and invasion potential of the round goby

European round gobies (Neogobius melanostomus) are displacing several important native North American fish species. Controlling their invasion is contingent on understanding their swimming inclination and potential. We assessed goby swimming inclination by recording activity in a 2 m flume over a ~24 h period, and swimming potential using a critical swimming (U _crit) test, as well as burst tests in still and flowing water. When given the choice to move, gobies covered as much as 14 m/h, with a slight bias towards nocturnal activity and an overall upstream preference. When confined and coerced to perform a U _crit test, they burst-and-held to achieve 35.5 ± 1.1 cm/s. Thirty minutes following U _crit, they were able to burst-and-coast in a sprint test to almost twice this speed. In still water, they exhibited startle bursts of up to 163 cm/s. We provide a swimming endurance model that indicates flow rates would need to be >125 cm/s to prevent upstream movement, and free of refuge areas in which to recover. The current study shows that the round goby is a surprisingly powerful swimmer with the capacity to continue its invasion should hydrologic control be absent.     

Critical swimming speed of yellow‐ and silver‐phase European eel (Anguilla anguilla,L.)

The prime objective of this study was to evaluate differences between the swimming performance of two distinct life stages of European eels. The critical swimming speed (U_crit) of 29 yellow‐ and 33 silver‐phase eels was evaluated in a swim tunnel. Silver‐phase eels showed a better swimming performance (U_crit = 0.66 ms^?1) than yellow individuals (U_crit = 0.43 ms^?1). Male and female silver eels reached an identical U_crit despite their different sizes, which may be a strategy to increase the synchronization of arrival at the spawning grounds.     

Divergence in physiological factors affecting swimming performance between anadromous and resident populations of brook charr Salvelinus fontinalis

In this study, an anadromous strain (L) and a freshwater‐resident (R) strain of brook charr Salvelinus fontinalis as well as their reciprocal hybrids, were reared in a common environment and submitted to swimming tests combined with salinity challenges. The critical swimming speeds (U_crit) of the different crosses were measured in both fresh (FW) and salt water (SW) and the variations in several physiological traits (osmotic, energetic and metabolic capacities) that are predicted to influence swimming performance were documented. Anadromous and resident fish reached the same U_crit in both FW and SW, with U_crit being 14% lower in SW compared with FW. The strains, however, seemed to use different underlying strategies: the anadromous strain relied on its streamlined body shape and higher osmoregulatory capacity, while the resident strain had greater citrate synthase (FW) and lactate dehydrogenase (FW, SW) capacity and either greater initial stores or more efficient use of liver (FW, SW) and muscle (FW) glycogen during exercise. Compared with R_♀L_♂ hybrids, L_♀R_♂ hybrids had a 20% lower swimming speed, which was associated with a 24% smaller cardio‐somatic index and higher physiological costs. Thus swimming performance depends on cross direction (i.e. which parental line was used as dam or sire). The study thus suggests that divergent physiological factors between anadromous and resident S. fontinalis may result in similar swimming capacities that are adapted to their respective lifestyles.     

Ontogenetic changes in the critical swimming speed of Gadus morhua (Atlantic cod) and Myoxocephalus scorpius (shorthorn sculpin) larvae and the role of temperature

Most studies on behavioural contributions to dispersal and recruitment during early life history stages of fishes have focused on coral reef species. For cold ocean environments, high variation in seasonal temperature and development times suggest that parallel studies on active behaviour are needed for cold-water species. Thus, we examined the critical swimming speed (U_crit) of marine fish larvae from 2 contrasting species: Gadus morhua (Atlantic cod) and Myoxocephalus scorpius (shorthorn sculpin), a pelagic and bottom spawner respectively. Within-species comparisons showed that sculpin reared at 6 °C had lower initial U_crit values, but a faster U_crit increase through development compared with 3 °C conspecifics, ultimately resulting in faster critical swimming speeds at metamorphosis (10.5 vs. 9.1 cm·s^− 1). In contrast, although cod larvae reared at 10 °C were faster swimmers at first feeding than 6 °C fish, temperature differences were absent after the first week. These results show that temperature influences the trajectory of larval critical swimming speed development, but that the relationship is species-specific. Although 6 °C sculpin and cod of similar length had equivalent U_crit values, the smaller size of cod at hatch (5.3 vs. 10.8 mm for sculpin) resulted in much lower age-specific U_crit values for cod. These data have significant implications for how swimming activity of the two species might affect dispersal, particularly in the first few weeks post-hatch. Overall, our data suggest that temperature during larval development influences the swimming capacity of cold-water marine fishes, and has important ramifications for biophysical models of dispersal.     

Thermal acclimation to 4 or 10°C imparts minimal benefit on swimming performance in Atlantic cod (Gadus morhua L.)

Thermal acclimation is frequently cited as a means by which ectothermic animals improve their Darwinian fitness, i.e. the beneficial acclimation hypothesis. As the critical swimming speed (U _crit) test is often used as a proxy measure of fitness, we acclimated Atlantic cod (Gadus morhua) to 4 and 10°C and then assessed their U _crit swimming performance at their respective acclimation temperatures and during acute temperature reversal. Because phenotypic differences exist between different populations of cod, we undertook these experiments in two different populations, North Sea cod and North East Arctic cod. Acclimation to 4 or 10°C had a minimal effect on swimming performance or U _crit, however test temperature did, with all groups having a 10–17% higher U _crit at 10°C. The swimming efficiency was significantly lower in all groups at 4°C arguably due to the compression of the muscle fibre recruitment order. This also led to a reduction in the duration of “kick and glide” swimming at 4°C. No significant differences were seen between the two populations in any of the measured parameters, due possibly to the extended acclimation period. Our data indicate that acclimation imparts little benefit on U _crit swimming test in Atlantic cod. Further efforts need to identify the functional consequences of the long-term thermal acclimation process.     

The effects of feeding on the swimming performance and metabolic response of juvenile southern catfish,Silurus meridionalis,acclimated at different temperatures

To test whether the effects of feeding on swimming performance vary with acclimation temperature in juvenile southern catfish (Silurus meridionalis), we investigated the specific dynamic action (SDA) and swimming performance of fasting and feeding fish at acclimation temperatures of 15, 21, 27, and 33 °C. Feeding had no effect on the critical swimming speeding (U_crit) of fish acclimated at 15 °C (p = 0.66), whereas it elicited a 12.04, 18.70, and 20.98% decrease in U_crit for fish acclimated at 21, 27 and 33 °C, respectively (p < 0.05). Both the maximal postprandial oxygen consumption rate (VO_2peak) and the active metabolic rate (VO_2active, maximal aerobic sustainable metabolic rate of fasting fish) increased significantly with temperature (p < 0.05). The postprandial maximum oxygen consumption rates during swimming (VO_2max) were higher than the VO_2active of fasting fish at all temperature groups (p < 0.05). The VO_2max increased with increasing temperature, but the relative residual metabolic scope (VO_2max? VO_2peak) during swimming decreased with increasing in temperature. The present study showed that the impairment of postprandial swimming performance increased with increasing temperature due to the unparalleled changes in the catfish's central cardio-respiratory, peripheral digestive and locomotory capacities. The different metabolic strategies of juvenile southern catfish at different temperatures may relate to changes in oxygen demand, imbalances in ion fluxes and dissolved oxygen levels with changes in temperature.     

The critical swimming speed of Iberian barbel Barbus bocagei in relation to size and sex

The swimming capacity of Barbus bocagei was measured with the critical swimming speed (U_crit) standard test in a modified Bla?ka‐type swim tunnel. Sixty B. bocagei were tested and they exhibited a mean ±s .d . U_crit of 0·81 ± 0·11 m s^?1 or 3·1 ± 0·86 total lengths per second (L_T s^?1). Sex had no effect on U_crit but significant differences were found between the swimming performance of fish with distinct sizes.     

Effects of age and size on critical swimming speed of juvenile Chinese sturgeon Acipenser sinensis at seasonal temperatures

Changes in the critical swimming speed (U_crit, cm s^?1) with ontogeny of 2·5–12·5 month‐old juvenile anadromous Chinese sturgeon Acipenser sinesis were measured in a modified Blazka‐type swimming tunnel. The absolute U_crit increased with length, mass and age; the relative U^′_crit (body lengths, s^?1), however, decreased. Juvenile A. sinesis did not display a parr–smolt transformation at the length or age threshold to tolerate full‐strength seawater.     

Swimming performance of two Iberian cyprinids: the Tagus nase Pseudochondrostoma polylepis (Steindachner, 1864) and the bordallo Squalius carolitertii (Doadrio, 1988)

The main purpose of this study was to gather swimming performance information for two endemic cyprinids of the Iberian Peninsula to contribute to the optimization of fish ways. Critical swimming speed (U_crit) was determined for the Tagus nase Pseudochondrostoma polylepis (Steindachner, 1864) and for the bordallo Squalius carolitertii (Doadrio, 1988) in a swimming tunnel. From a total of 80 P. polylepis tested, the mean (± SD) U_crit observed was 0.78 ± 0.15 ms^?1 (c. 3.74 ± 0.93 BL s^?1); the 68 S. carolitertii tested presented an U_crit of 0.54 ± 0.1 ms^?1 (c. 4.43 ± 0.74 BL s^?1). Significant interspecific differences were found between the U_crit of the tested cyprinids. Intraspecific comparisons between the U_crit and the variables of size, sex, condition factor and gonado‐somatic index were also made. No sex‐or gonad maturation‐related differences between the U_crit were identified, but the robust P. polylepis were found to be stronger swimmers. Water velocities in fish ways for P. polylepis and S. carolitertii should aim, on average, for lower than 0.7 and 0.5 ms^?1, respectively.     

Gastrointestinal blood flow and postprandial metabolism in swimming sea bass Dicentrarchus labrax

In trout and salmon, the metabolic costs of exercise and feeding are additive, which would suggest that gastrointestinal blood flow during exercise is maintained to preserve digestive and absorptive processes related to the specific dynamic action (SDA) of food. However, in most published studies, gastrointestinal blood flow drops during swimming, hypoxia, and general stress. To test whether gastrointestinal blood flow is spared during exercise after feeding, sea bass were instrumented with flow probes to measure cardiac output and celiacomesenteric blood flow while swimming in a respirometer before and after feeding. Swimming at 2 body lengths per second (bl s(-1)) increased metabolic rate considerably more than did feeding (208% vs. 32% increase, respectively, relative to resting), and a similar pattern was observed for cardiac output. In unfed fish, resting gastrointestinal blood flow was 13.8+/-0.5 mL min(-1) kg(-1). After feeding, resting gastrointestinal blood flow increased by 82% but then decreased progressively with increasing swimming speeds. At 2 bl s(-1), gastrointestinal blood flow in fed fish was not significantly different compared with that in unfed swimming fish, and, therefore, the data do not support the gastrointestinal sparing hypothesis. The magnitude of the SDA was maintained despite the decrease in gastrointestinal blood flow and the consequent reduction in oxygen supply to the gut. An estimate of maximal oxygen flow to the gastrointestinal tract after feeding yielded 2.6 mmol O(2) h(-1) kg(-1), but this amount is not able to cover the oxygen demand of 3.16 mmol O(2) h(-1) kg(-1). Therefore, the SDA must reflect metabolic processes in tissues other than those directly perfused by the celiacomesenteric artery.