On the interaction of turbulent flow and feeding behaviour of juvenile Atlantic Salmon (Salmo salar)

Anthropogenic activities can dramatically modify the riverine habitat of juvenile Atlantic salmon ( Salmo salar ). In the perspective of protecting and restoring the fluvial habitat, bioenergetic models are often used to estimate fish habitat quality. These models determine the habitat quality as the ratio between the energetic gains (food) and costs (growth, metabolism) of a fish. The energetic costs of swimming in a river are generally estimated using the average flow velocity without consideration of the effect of turbulence. Juvenile Atlantic salmon (JAS) live in rivers characterized by intense velocity fluctuations, which are often described as a succession of high‐ and low‐speed flow regions. These flow structures are likely to affect the JAS activity that consists of long periods of sit‐and‐wait at the top of a protuberant rock interrupted by short bursting motions to capture drifting food particles. To minimize the energetic costs, it is hypothesized that JAS use low‐speed flow regions to initiate and undertake their feeding motions.  To improve bioenergetic modelling, this study aimed at analyzing the relation between turbulent flow structures and the feeding behaviour of JAS in a natural gravel‐bed river. We filmed eight JAS during 30 min with a submersible video camera while simultaneously measuring velocity fluctuations close to the fish in the St. Marguerite River, Quebec, Canada. Our results show that the proportion of time used for feeding motions decreases with increasing turbulent intensity and mean flow velocity; and that JAS do not seem to prefer low‐speed flow regions to initiate their feeding motions.     

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.     

鲢幼鱼通过水流速度障碍的模拟

鱼类能否通过水流速度障碍直接影响过鱼设施的过鱼效果。利用计算机技术,综合水力因素、鱼类行为、地理特征及环境因子,展开鱼类通过水流速度障碍的模拟,有助于过鱼设施的优化设计。以国外涵洞式鱼道模拟软件Fish Xing为切入点,结合主要模块和关键因子,对我国特有鱼类鲢幼鱼进行模拟,得到鲢通过不同水流速度障碍的成功率;对比鲢在物理模型中的游泳表现,从模型主要模块和影响鱼类游泳表现的关键因子角度,分析影响鱼类通过水流速度障碍模拟的因素。结果表明,Fish Xing软件不能精确模拟鲢通过水流速度障碍的表现。分析表明,该软件在地理要素、管道特征和水力信息等参数方面具备独特的优势,但对我国鱼类有一定局限性,主要体现在鱼类的生物学信息如鱼类游泳特征等方面存在不足;进行鱼过障碍的模拟需要深入研究目标鱼类的生理特征、游泳能力及其与水力环境因子的响应关系。     

五种淡水鱼类幼鱼游泳能力的比较

为了探讨栖息于不同生境中鱼类的游泳能力和偏好游泳速度及其生理机制,本研究以中华倒刺鲃(Spinibarbus sinensis)、异育银鲫(Carassius auratus gibelio)、岩原鲤(Procypris rabaudi)、青鱼(Mylopharyngodon piceus)和胭脂鱼(Myxocryprinus asiaticus)5种鱼的幼鱼为对象,在(25±1)℃条件下测定了5种鱼类的标准代谢率(SMR)、最大代谢率(MMR)、有氧代谢范围(MS)、临界游泳速度(Ucrit)、最大匀加速游泳速度(Ucat)和偏好游泳速度(Upref)。结果发现:5种实验鱼中,中华倒刺鲃的游泳能力最强,游泳能力较差的为青鱼和胭脂鱼;5种鱼之间的代谢和游泳能力差异显著,其偏好游泳速度主要集中在(10~24.5cm·s^-1)区域。研究表明,鱼类游泳能力的种间差异可能主要由心鳃系统相关的呼吸能力和体型相关的游泳效率所决定。本研究提供的有关鱼类游泳能力、偏好游泳速度等资料对于鱼道设计等有一定的参考价值。     

The effect of substratum type on aspects of swimming performance and behaviour in shortnose sturgeon Acipenser brevirostrum

The swimming performance and associated swimming behaviour (i.e. substratum‐skimming, station‐holding and free swimming) were assessed in shortnose sturgeon Acipenser brevirostrum during critical swimming and endurance swimming tests over a rough and a smooth substratum. It was hypothesized that the addition of a rough substratum in the swimming flume may provide a surface for the A. brevirostrum to grip and offer an energetic advantage. Substratum type did not affect the critical swimming performance, but A. brevirostrum consistently performed more bottom behaviours (i.e. substratum‐skimming and station‐holding) while on a smooth substratum. Acipenser brevirostrum had little contact with the rough substratum until the velocity was >1 body length s^?1. Endurance swimming time was significantly lower for A. brevirostrum over the rough bottom at the highest velocity (30 cm s^?1) which may be attributed to the observed increase in free swimming and decrease in bottom behaviours. During endurance swimming, the rough substratum was mainly used at intermediate velocities, suggesting that there may be a stability cost associated with being in contact with the rough substratum at certain velocities.     

A split decision: the impact of substrate type on the swimming behaviour,substrate preference and UCrit of juvenile shortnose sturgeon (<Emphasis Type="Italic">Acipenser brevirostrum</Emphasis>)

Critical swimming speed (UCrit) is a standard test to measure sustained swimming capabilities of fish species, however, much of this research is focused on pelagic fish or popular game fish. Recently, more research is emerging on the swimming capabilities of sturgeons, mainly due to their conservation status. Substrate preference has been examined in sturgeons, however, few studies have investigated whether sturgeon would select for a particular substrate (smooth or pebble) when provided a choice under high-flow conditions, as irregular shaped bottom substrates may provide an energetic advantage. Critical swimming tests were performed in a linear flume to evaluate whether substrate (smooth, pebble, pebble on the left side of the flume/smooth on the right side of the flume, and pebble on the right side of the flume/smooth on the left side of the flume) would affect UCrit of juvenile shortnose sturgeon (Acipenser brevirostrum), and if the sturgeon would select for a particular substrate at different speeds. All swimming experiments were video recorded and subsequently reviewed to determine the amount of time individual sturgeon spent in particular sections of the flume. Overall, there was no clear preference for a particular substrate and substrate configuration did not affect UCrit. These results may be attributed to small sample sizes, and the small pebble sizes used in relation to the sturgeon’s body size may not elicit an energetic advantage.     

Hydraulic aspects of pool-weir fishways as ecologically friendly water structure

Today, natural rivers are increasingly fragmented by human-made obstacles such as dams. Due to these structures fish populations in natural rivers rapidly diminish. In this context, fishways are a useful hydraulic structure in the creation of ecosystem for fish migration. Three dimensional mean flow and turbulence structure of pool-weir fishways were experimentally explored. During the experiments, three different notch sizes were applied while the size of the orifice was kept constant. Two acoustic Doppler velocimeters were employed throughout the velocity measurements. Three-dimensional mean velocity and normalized turbulent kinetic energy patterns in the pool were experimentally analyzed considering the swimming ability of different fish species to check whether the given design conditions provide suitable flow patterns. Based on the data, a linear relationship between the parameters “the discharge” and “the average depth in a pool” was generated. An equation was derived which gives the “energy dissipation rate per unit pool volume” in terms of the parameters “geometrical characteristics of the fishway”, “head difference between pools”, “slope”, and “acceleration due to gravity”. The discharge ratios between “flow through orifice” and “flow over notch” were expressed based on the data.     

基于雅砻江两种裂腹鱼游泳能力的鱼道设计

为探究雅砻江两种裂腹鱼的游泳能力,给过鱼设施设计和鱼类游泳行为学研究提供基础参数,本研究采用递增流速法对长丝裂腹鱼、齐口裂腹鱼的感应流速、临界游泳速度、突进游泳速度进行测试,采用固定流速法对长丝裂腹鱼的耐久游泳速度进行测试。结果表明: 长丝裂腹鱼与齐口裂腹鱼的感应流速随着体长的增加均出现了先增加后平稳的趋势,但最大感应流速均小于0.2 m·s^-1;长丝裂腹鱼的临界游泳速度与突进游泳速度分别为(0.81±0.20)和(1.49±0.26) m·s^-1,相对临界游泳速度为(4.90±1.73) BL·s^-1,相对突进游泳速度为(9.77±1.72) BL·s^-1(BL为体长);齐口裂腹鱼的临界游泳速度与突进游泳速度分别为(0.73±0.24)和(1.17±0.39) m·s^-1,相对临界游泳速度为(6.88±2.82) BL·s^-1,相对突进游泳速度为(11.75±2.77) BL·s^-1。耐久测试发现,随着流速增加(0.7～1.5) m·s^-1,长丝裂腹鱼持续游泳时间与水流速度呈负相关,疲劳时间(T)与水流速度(V)的关系可以拟合为lgT=-2.52V+5.59,预测鱼道长度(d)与鱼道内可通过的最大平均水流速度(V_fmax)的关系式为V_{f max}=-0.17lnd+1.74。根据试验结果,当以长丝裂腹鱼和齐口裂腹鱼为主要过鱼对象时,建议鱼道内最小水流速度应大于0.2 m·s^-1,进口及竖缝处水流速度为0.73～1.67 m·s^-1,休息池主流水流速度为0.2～0.7 m·s^-1。     

Behaviour and Locomotor Activity of a Migratory Catostomid during Fishway Passage

Fishways have been developed to restore longitudinal connectivity in rivers. Despite their potential for aiding fish passage, fishways may represent a source of significant energetic expenditure for fish as they are highly turbulent environments. Nonetheless, our understanding of the physiological mechanisms underpinning fishway passage of fish is still limited. We examined swimming behaviour and activity of silver redhorse (Moxostoma anisurum) during its upriver spawning migration in a vertical slot fishway. We used an accelerometer-derived instantaneous activity metric (overall dynamic body acceleration) to estimate location-specific swimming activity. Silver redhorse demonstrated progressive increases in activity during upstream fishway passage. Moreover, location-specific passage duration decreased with an increasing number of passage attempts. Turning basins and the most upstream basin were found to delay fish passage. No relationship was found between basin-specific passage duration and activity and the respective values from previous basins. The results demonstrate that successful fishway passage requires periods of high activity. The resultant energetic expenditure may affect fitness, foraging behaviour and increase susceptibility to predation, compromising population sustainability. This study highlights the need to understand the physiological mechanisms underpinning fishway passage to improve future designs and interpretation of biological evaluations.     

Effects of fish size, time‐to‐fatigue and turbulence on swimming performance: a case study of Galaxias maculatus

A simple relationship for the inanga Galaxias maculatus swimming velocity is suggested and tested in low and high turbulence channels. The relationship connects the swimming velocity with fish Reynolds and Froude numbers and can be used in both ecological analysis ( e.g . habitat requirements) and management strategies ( e.g . fishways design). Contrary to some previous studies and intuition, effects of turbulence on swimming performance appeared to be negligible. The most likely explanation for this result relates to mechanics of fish–turbulence interactions that may be dependent on both the turbulence scales and energy. The data suggest that future studies of turbulence effects on fish behaviour should involve, in addition to turbulence energetics, consideration of fish dimensions in relation to the spectrum of turbulence scales.     

Swimming efficiency and the influence of morphology on swimming costs in fishes

Swimming performance is considered a main character determining survival in many aquatic animals. Body morphology highly influences the energetic costs and efficiency of swimming and sets general limits on a species capacity to use habitats and foods. For two cyprinid fishes with different morphological characteristics, carp (Cyprinus carpio L.) and roach (Rutilus rutilus (L.)), optimum swimming speeds (U _mc) as well as total and net costs of transport (COT, NCOT) were determined to evaluate differences in their swimming efficiency. Costs of transport and optimum speeds proved to be allometric functions of fish mass. NCOT was higher but U _mc was lower in carp, indicating a lower swimming efficiency compared to roach. The differences in swimming costs are attributed to the different ecological demands of the species and could partly be explained by their morphological characteristics. Body fineness ratios were used to quantify the influence of body shape on activity costs. This factor proved to be significantly different between the species, indicating a better streamlining in roach with values closer to the optimum body form for efficient swimming. Net swimming costs were directly related to fish morphology.     

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.     

Thermal acclimation effects differ between voluntary, maximum, and critical swimming velocities in two cyprinid fishes

Temperature acclimation may be a critical component of the locomotor physiology and ecology of ectothermic animals, particularly those living in eurythermal environments. Several studies of fish report striking acclimation of biochemical and kinetic properties in isolated muscle. However, the relatively few studies of whole-animal performance report variable acclimation responses. We test the hypothesis that different types of whole-animal locomotion will respond differently to temperature acclimation, probably due to divergent physiological bases of locomotion. We studied two cyprinid fishes, tinfoil barbs (Puntius schwanenfeldii) and river barbels (Barbus barbus). Study fish were acclimated to either cold or warm temperatures for at least 6 wk and then assayed at four test temperatures for three types of swimming performance. We measured voluntary swimming velocity to estimate routine locomotor behavior, maximum fast start velocity to estimate anaerobic capacity, and critical swimming velocity to estimate primarily aerobic capacity. All three performance measures showed some acute thermal dependence, generally a positive correlation between swimming speed and test temperature. However, each performance measure responded quite differently to acclimation. Critical speeds acclimated strongly, maximum speeds not at all, and voluntary speeds uniquely in each species. Thus we conclude that long-term temperature exposure can have very different consequences for different types of locomotion, consistent with our hypothesis. The data also address previous hypotheses that predict that polyploid and eurythermal fish will have greater acclimation abilities than other fish, due to increased genetic flexibility and ecological selection, respectively. Our results conflict with these predictions. River barbels are eurythermal polyploids and tinfoil barbs stenothermal diploids, yet voluntary swimming acclimated strongly in tinfoil barbs and minimally in river barbels, and acclimation was otherwise comparable.     

Empirical analysis of the influence of swimming pattern on the net energetic cost of swimming in fishes

We tested the hypothesis that the energetics of swimming in a flume accurately represent the costs of various spontaneous movements using empirical relationships between fish swimming costs, weight, and speed for three swimming patterns: (1) 'forced swimming' corresponded to movements adopted by fish forced to swim against a unidirectional current of constant velocity; (2) 'directed swimming' was defined as quasi-rectilinear movements executed at relatively constant speeds in a stationary body of water and (3) 'routine swimming' was characterized by marked changes in swimming direction and speed. Weight and speed explained between 76% (routine swimming) and 80% (forced swimming) of net swimming cost variability. Net costs associated with different swimming patterns were compared using ratios of model predictions (swimming cost ratio; SCR) for various weight and speed combinations. Routine swimming was the most expensive swimming pattern (SCR for routine and forced swimming =6.4 to 14.0) followed by directed (SCR for directed and forced swimming =0.9 to 2.8), and forced swimming. The magnitude of the difference between the net costs of forced and spontaneous swimming increases with movement complexity and decreases as fish weight increases.     

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.     

Water velocity shapes fish movement behavior

Stream and river ecosystems present fluvial fishes with a dynamic energy landscape because moving water generates heterogeneous flow fields that are rarely static in space and time. Fish movement behavior should be consistent with conserving energy in these dynamic flowing environments, but little evidence supporting this hypothesis exists. Here, we tested experimentally whether three general movement behaviors—against the current, with the current, or holding position (i.e., staying in one position and location)—were performed in a way consistent with minimizing the cost of swimming in a heterogeneous flow field. We tested the effects of water velocity on movement behavior across three age classes (0, 1, and 5 years) of two different fluvial specialist fishes, the pallid sturgeon (Scaphirhynchus albus) and shovelnose sturgeon (Scaphirhynchus platorynchus). Individuals from the three age classes were exposed to a continuous and dynamic velocity field ranging from 0.02 to 0.53 m s⁻¹, which represented natural benthic flow regimes occupied by these species in rivers. Both sturgeon species exhibited the same pattern with regard to their tendency to hold position, move upstream, or move downstream. Moving downstream was positively associated with velocity for all age groups. Moving upstream was inversely related to velocity for young fish, but as the fish aged, moving upstream was not related to water velocity. The oldest fish (age 5) moved upstream more frequently compared to the younger age classes. Holding position within a water current was the most frequent behavior and occurred with similar probability across the range of experimental velocity for youngest fish (age 0), but was inversely related to velocity in older fish. Our experiment across age classes suggests that the suite of swimming behaviors exhibited by fluvial specialists might have evolved to mitigate the energetic costs of complex energy landscapes generated by moving water to ultimately maximize net energy gain.     

Postprandial intestinal blood flow, metabolic rates, and exercise in Chinook salmon (Oncorhynchus tshawytscha)

Following a relatively large meal (2% body mass of dry pellets), intestinal blood flow in chinook salmon (Oncorhynchus tshawytscha) increased significantly, up to 81%, between 14 and 29 h postprandially. Also, 15 h postprandially, oxygen consumption (M(2)) was elevated by 128% compared with a measurement of routine M(2) made after 1 wk of fasting. The postprandial increase in MO(2) (the heat increment) was 33 micromol O(2) min(-1) kg(-1). Because intestinal blood flow is known to decrease during swimming activity in fish, we therefore tested the hypothesis that swimming fish would have to make a trade-off between maximum swimming activity and digestive activity by comparing the swimming performance and metabolic rates of fed and fasted chinook salmon. As expected, MO(2) increased exponentially with swimming velocity in both fed and fasted fish. Moreover, the heat increment was irreducible during swimming, such that MO(2) remained approximately 39 micromol O(2) min(-1) kg(-1) higher in fed fish than in fasted fish at all comparable swimming speeds. However, maximum M dot o2 was unaffected by feeding and was identical in both fed and fasted fish (approximately 250 micromol O(2) min(-1) kg(-1)), and, as a result, the critical swimming speed (U(crit)) was 9% lower in the fed fish. Three days after the fish were fed and digestion was completed, MO(2) and U(crit) were not significantly different from those measured in fasted fish. The ability of salmonids to maintain feeding metabolism during prolonged swimming performance is discussed, and it is suggested that reduced swimming performance may be due to postprandial sparing of intestinal blood to support digestion, thereby limiting the allocation of blood flow to locomotory muscles.     

Numerical flow simulation of pool-type fishways: new ways with well-known tools

Fish passage structures are built to restore the connectivity of rivers and allow the migration of aquatic fauna. In order to assess the functioning of a pool-type fishway, it is necessary, inter alia, to possess detailed knowledge of its flow structure, since observations of fishways, and in particular of the visible water surface, can only provide a rough idea of the actual conditions inside the pools. Numerical simulation has been used for many years to support engineering sciences. Especially, the modeling of flow processes in hydraulic machines can, on the one hand, help avoid major problems during the design stage of fish passage structures and, on the other, improve the structure’s hydraulic performance. To this end, two diploma theses within the framework of a research project of the local energy supplier Energie Baden-Württemberg AG (EnBW) employed modeling tools for 3D flow simulation, primarily for pool-slot fishways (PSF), and for traditional vertical slot fishways (VSF). Guest editors: R. L. Welcomme & G. Marmulla Hydropower, Flood Control and Water Abstraction: Implications for Fish and Fisheries     

Models for tuna school formation

Schooling behavior is a challenging topic in the context of animal aggregation. It is also of economic importance for the estimation and conservation of stock sizes. An individual based movement model will be developed, taking into account energetic advantages of schooling. This model is a cellular automaton with a hexagonal grid. The latter considers the geometry of a school, where fish swim in a diamond-shape configuration in order to take advantage of the velocity, induced by the tail strokes of preceding fish. Furthermore, knowing the induced velocity field makes it possible to consider the energetic needs of fish swimming in that school and to describe the break up of schools due to oxygen depletion. This allows us to estimate maximum school sizes.     

基于物理模型实验的光倒刺鲃生态行为学研究

建立了基于天然河道的物理模型,通过控制实验研究水环境因子对鱼类行为的影响。文中选取了金沙江下游2.6 km河段建立物理模型,以南方水系的经济鱼类光倒刺鲃为研究对象,进行了鱼的行为对底质和流速的响应实验。分析得出,光倒刺鲃对砂卵石底质的选择明显大于其它底质且差异极其显著(P<0.01),光倒刺鲃2龄幼鱼期的喜好流速范围为0.3-0.6 m/s;研究同时发现水流紊动强度对光倒刺鲃行为具有重要影响。该研究结果可以为光倒刺鲃栖息地模型提供参考。