Light-oriented swimming of schooling fish in continuous channels

Synopsis Fish groups were tested both in a circular and in a figure eight-shaped channel. In both cases fish showed a long lasting, constant direction swimming provided that illumination was maintained at a constant angle around the channel. In the circular channel, fish did not reverse direction, as would be expected, when light angle was shifted from one side to the other in the channel. However, direction reversals did occur when these illumination shifts were performed on the eight-shaped channel. We suggest that constant-oriented swimming reflects a sun-compass oriented behavior, but swimming at a constant angle in the circular channel produces an irreversible disarrangement of the inertial-orientation system, which does not occur in the eight-shaped channel due to the geometrical relationship between the light and the shape of the channel.     

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.     

Schooling behaviour and retinomotor response of juvenile Pacific bluefin tuna Thunnus orientalis under different light intensities

Schooling behaviour and histological retinal light adaptation in juvenile Pacific bluefin tuna Thunnus orientalis were examined under various light intensities to determine the effect of light intensity on behaviour. After monitoring the schooling behaviour of juveniles 35–36 and 45–46 days post hatching, schooling variables such as nearest neighbour distance and separation swimming index were measured under different light intensities. Furthermore, retinal indices of light adaptation were investigated histologically for each experimental light intensity. Under intensities >5 lx, schooling variables in the two juvenile growth stages were nearly constant, allowing schooling. In contrast, the schooling variables indicated that the fish gradually swam more widely and randomly with decreasing light intensities <5 lx. The retinal indices also showed a shift from light adaptation to dark adaptation at light levels <5 lx. From 5 to 0·01 lx, retinal adaptation and fish schooling behaviour changed with light intensity. These data suggest that the schooling behaviour of juvenile Pacific bluefin tuna is greatly affected by retinal adaptation.     

Alterations to the swimming performance of carp, Cyprinus carpio, as a result of temperature acclimation

Groups of common carp were acclimated to either 10°C or 28°C for 6 weeks. Fish were then exercised at 10°C or 20°C, and the critical swimming speed (fatigue velocity) was measured. At 10°C, cold-acclimated carp were capable of significantly higher swimming speeds. When exercised at 20°C. however, the situation was reversed, and warm-acclimated carp exhibited improved swimming ability. These results provide direct evidence that acclimation of the contractile proteins is beneficial across a wide temperature range. Following acclimation to low environmental temperatures the critical swimming speed exhibited a Q₁₀ of only 1.1 for the temperature range 10–20°C. compared to a value of 2.9 for fish acclimated to the higher temperature.     

Reactions in individual fish to strobe light. Field and aquarium experiments performed on whitefish (Coregonus lavaretus)

This study describes how individual whitefish Coregonus lavaretusreact to strobe light. Field experiments were performed in a net enclosure on fish tagged with ultrasonic transmitters. A strobe light array was switched on near the tagged fish. The fish moved away from the light and increased their swimming speed. Aquarium experiments under controlled conditions were carried out in rearing tanks at Saimaa Fisheries and Aquaculture Station in Finland. A strobe light was directed from the side of the basin just ahead of, directly at, and behind the fish at a close range. In the first two cases fish responded by a distinct turn and a change in swimming direction away from the light. The fish did not change its swimming direction when light was aimed from behind. It is concluded that strobe light may be used to prevent fish from swimming into a specific area. Implications for development of new fishing equipment and research concerning fishes in areas with water power stations is briefly discussed.     

Behavioural studies on the lesser sandeel Ammodytes marinus (Raitt) III. The effect of temperature on activity and the environmental control of the annual cycle of activity

The behaviour of the lesser sandeel, Ammodytes marinus (Raitt), has been investigated at 5, 10 and 15° C, using a photographic method of recording activity. The activity patterns at 10 and 15° C were very similar, there being a high level of swimming activity during the light period, which fell to a low level at 5° C. It was also lower at 10° C at the end of the experiment than at the beginning and it is suggested that this might have been due to an increase in the fat contents of the fish. The feeding rate of the fish was measured and showed a Q ₁₀ of 2.08 for the temperature range 5–15° C. The annual cycle of activity of A. marinus is discussed in relation to seasonal changes in food availability, light and temperature, and in the fat content of the fish. It is concluded that after spawning in the December–January period the fish remain buried in the sand until April, because of the limiting effect on swimming and feeding activity of the environmental factors in the intervening period. The proportion of fish available for capture at the start of the fishery in April is related mainly to temperature, but food (as measured by numbers of copepods) light intensity and photoperiod are by then increasing rapidly. After July the fishery ceases and it is thought that this is because the fish have entered an overwintering stage, during which they remain buried in the sand. This phase is also thought to be associated with the maturation of the gonads in readiness for the winter spawning. The factors causing the fish to enter this stage are as yet undetermined but may be related to the attainment of a certain level of fat content.     

The ontogeny of schooling behaviour in the striped jack

The ontogeny of schooling behaviour was investigated in laboratory-reared striped jack Pseudocaranx dentex larvae and juveniles. Mean separation angle between nearest individuals averaged 78 and 82° in 10- and 12-mm fish, respectively, and decreased to 57° in 16-mm fish. In addition, interindividual distance decreased from three times body length in 12-mm fish to twice the body length in 16-mm individuals; this value continued to decrease gradually to 1·3 at 19 mm to 0·79 at 30 mm. Mutual attraction due to visual recognition was observed in 12-mm fish and increased with total length. The optokinetic response ( R_o , the ability of a fish to keep station with moving patterns) first appeared in 4–6-mm larvae. In spite of the traditional emphasis of the importance of R_o in schooling behaviour, the onset of R_o was insufficient to result in school formation. The emergence of mutual attraction seems essential for the ontogeny of schooling.     

Behavioural studies on the lesser sandeel Ammodytes marinus (Raitt) II. The effect of light intensity on activity

The behaviour of the lesser sandeel, Ammodytes marinus (Raitt), has been investigated at light intensities of 1, 10, 100 and 1000 lux, using a photographic method of recording activity. The level of swimming activity was high at 1000 and 100 lux, declining to a very low level at 1 lux. It was concluded that this was due to the limiting effect of light on feeding. The threshold light intensity for swimming activity in the tank was estimated as being approximately 20 lux but it was considered that in the area of the sandeel fishing grounds the threshold might be higher than this, in the region of 100 lux. The number of hours light per day above 20 and 100 lux at a depth of 15 m in the area of the Outer Dowsing sandbank (53°30'N, 01°00'E) was estimated for the various months of the year. It was shown that during the winter the light intensity does not normally reach 100 lux and only exceeds 20 lux for a few hours each day. It is suggested that this could limit swimming activity and accessibility at this time of year. Measurements were made of the penetration of light into sand and it was concluded that fish which are buried might be able to detect light, possibly via the pineal gland.     

Circadian rhythms in juvenile american shad, Alosa sapidissima

Swimming activity (in cm s⁻¹) of a school (55 individuals) of young-of-the-year ( total length=110 mm) American shad, Alosa sapidissima , was determined under a variety of photoperiod conditions. These included a normal (ambient), a shifted, and constant-light day. Swimming activity was measured over 4-day periods. During normal days swimming speeds followed periods of about 24 h, with fast speeds (up to 45 cm s⁻¹) and schooling occurring during the photoperiod. Under dark conditions speeds were slower (8 cm s⁻¹) with most fish swimming as individuals. During a shifted day swimming speeds and schooling corresponded to the imposed day. Under constant light (equivalent to bright moonlight) no schooling was evident, and a constant, but slow, swimming speed was observed in each 24-h period. These shad demonstrated an exogenous rhythm with respect to the imposed day length. It is hypothesized that an endogenous circadian rhythm would only be of use to a fish required to hunt or chase its prey. Shad, being plankton feeders, do not chase prey and therefore can exhibit an exogenous circadian rhythm with no detrimental feeding results.     

Thermally induced phenotypic plasticity of swimming performance in European sea bass Dicentrarchus labrax juveniles

The vulnerability of embryonic and larval stages of European sea bass Dicentrarchus labrax to environmental temperature and the longer-term consequences for the early juveniles was demonstrated. This phenotypic plasticity was highlighted by subjecting D. labrax at 15·2 ± 0·3 or 20·0 ± 0·4° C (mean ± s . d .) up to metamorphosis and then at the same temperature (18·5 ± 0·7° C). After 4–6 weeks at the same temperature, the measurement of critical swimming speed at four exercise temperatures (15, 20, 25 and 28° C) showed a significantly higher swimming capacity in the fish initially reared at 15° C than for fish initially reared at 20° C. This performance was correlated with significant differences in the phenotype of red muscle. Thermally induced phenotypic plasticity was clearly demonstrated as an important mechanism controlling swimming performance in early juveniles of D. labrax .     

Is tilting behaviour at low swimming speeds unique to negatively buoyant fish? Observations on steelhead trout,Oncorhynchus mykiss,and bluegill,Lepomis macrochirus

When swimming at low speeds, steelhead trout and bluegill sunfish tilted the body at an angle to the mean swimming direction. Trout swam using continuous body/caudal fin undulation, with a positive (head-up) tilt angle ( 0 , degrees) that decreased with swimming speed ( u , cm s⁻¹) according to: 0 =(164±96).u^{(−1.14±0.41)} (regression coefficients; mean±2 s.e. ). Bluegill swimming gaits were more diverse and negative (head down) tilt angles were usual. Tilt angle was −3·0 ± 0.9° in pectoral fin swimming at speeds of approximately 0.2–1.7 body length s⁻¹ (Ls⁻¹; 3–24 cm s⁻¹), −4.5 ±2.6° during pectoral fin plus body/caudal fin swimming at 1·2–1·7 L s⁻¹ (17–24cm s⁻¹), and −5.0± 1.0° during continuous body/caudal fin swimming at 1.6 and 2.5 L s⁻¹ (22 and 35cm s⁻¹). At higher speeds, bluegill used burst-and-coast swimming for which the tilt angle was 0.1±0.6°. These observations suggest that tilting is a general phenomenon of low speed swimming at which stabilizers lose their effectiveness. Tilting is interpreted as an active compensatory mechanism associated with increased drag and concomitant increased propulsor velocities to provide better stabilizing forces. Increased drag associated with trimming also explains the well-known observation that the relationship between tail-beat frequency and swimming speed does not pass through the origin. Energy dissipated because of the drag increases at low swimming speeds is presumably smaller than that which would occur with unstable swimming.     

Laboratory experiments on the schooling and feeding behaviour of Trichogaster fasciatus (Pisces: Perciformes)

Laboratory experiments were performed on the schooling and feeding behaviour of Trichogaster fasciatus . There was a tendency on the part of individuals of similar size to school together; the instinct being stronger in younger specimens. Unavailability of similar-sized individuals led to schooling by members of different size-groups. Column feeding was exhibited by the fish. While feeding at the bottom, the fishes formed angles varying from 30°90°. Larger individuals formed a right angle more often than younger members of the species. Optimum degree of crowding seemed necessary for active feeding as confinement of single individual, or overcrowded condition, both resulted in a decline of feeding activity.     

Growth, pigmentation and activity of juvenile Japanese eels in relation to temperature and fish size

The growth and activity of juvenile Japanese eels Anguilla japonica in different pigmentation stages from the glass eel to the elver stage were studied in the laboratory at 15, 20 and 25° C. The growth and activity of the eels were significantly influenced by both temperature and fish size. Growth rate generally declined with increasing fish size, and fish were least active and experienced a low growth during the pigmenting stage at all temperatures. They were nocturnal and spent significantly more time moving (swimming, feeding and moving over the substratum) at 20 and 25° C than at 15° C at night within each pigmentation stage. Accordingly, they grew significantly faster at 20 and 25° C than at 15° C throughout the study. The development of pigmentation appeared to be dependant on water temperature but not on fish size. This study suggested that the growth and activity of juvenile Japanese eels were positively correlated, because fish were least active and grew slowest at low temperature (15° C) or during the pigmenting stage at all temperatures.     

School Formation and Concurrent Developmental Changes in Carangid Fish with Reference to Dietary Conditions

The ontogeny of schooling behaviour was studied in comparison to the development of sensory and swimming organs and taxis in carangid fish. Striped jack, Pseudocaranx dentex, larvae showed strong phototaxis at 3 days after hatching (3.5 mm in TL) when they developed pigmentation in the retina. Rheotaxis and optokinetic responses were apparent at 4.0-6.0 mm TL as larvae completed development of the basic structure of their eyes. A major inflection of allometric growth occurred at 9 mm, and fin ray formation was completed at these stages. Schooling behaviour, represented by one TL of inter-individual distance and parallel orientation, only appeared at 16 mm TL, and just prior to this behaviour, fish showed mutual attraction through vision at 12 mm TL. Canalization of buccal lateral lines was complete at 18 mm TL, whereas that of trunk lateral lines started at 23 mm TL and was complete at 30 mm TL. With these results, we assumed that critical factors of the ontogeny of schooling behaviour in carangid fish include not only the development of sensory or swimming organs, but also other factors such as development of the central nervous system. To show this, we reared another carangid species, the yellowtail Seriola quinqueradiata, with dietary depletion of DHA (docosahexaenoic acid), which is indispensable for the development of the central nervous system. Although DHA-free fish showed optokinetic response, they did not show schooling behaviour when they attained their schooling size. Tracer experiments using radioisotope labelled DHA showed that DHA is incorporated into the brain, spinal cord, and retina of juvenile fish. Under natural conditions, carangid fish larvae should intake enough DHA through diet to develop schooling behaviour, the fluctuation of dietary quality in zooplankton might therefore influence the development of indispensable antipredatory behaviour. Morphological changes of striped jack occurred in two steps; first at 9-12 mm (fin formation and inflection of allometric growth) and then second at 20-30 mm (scale and lateral line formation), and these changes corresponded with the development of schooling and recruitment to coastal waters, respectively. Since the onset of schooling is the first step of active antipredatory behaviour, we considered that 12 mm TL is the size at which they attain the juvenile period.     

A new type of metabolism chamber for the determination of active and postprandial metabolism offish, and consideration of results for coregonid and salmon juveniles

The optomotor reaction of juvenile Coregonus schinzipalea Val. et Cuv. and Salmo salar L. was utilized to develop a circular tube metabolism chamber to measure oxygen consumption and ammonia excretion as a function of swimming speed. The metabolism chamber with a constant water flow assured the maintenance of stable conditions. The unidirectional movement of fish was measured in a circular tube with a single narrowing. The relationships between the swimming speed and oxygen consumption or ammonia excretion described by exponential equations allowed the extrapolation towards the standard metabolism, i.e., zero swimming speed. For a juvenile coregonid (0.1–0.15 g individual weight, 2.6–2.8 cm total length) standard metabolism at 14° C was estimated as 0.65 mg0₂ g⁻¹ h⁻¹ and 17.3 μg N(NH₃)g⁻¹ h⁻¹, whereas for juvenile salmon (136mg individual weight) respective values at 22° C were 0.047mg0₂g⁻¹h⁻¹ and 0.61 μg N(NH₃)g⁻¹ h⁻¹. The feeding test with juvenile salmon was also performed in this circular chamber, and in both energy and nitrogen budgets after a meal the partitioning could be precisely attributed to standard metabolism, active metabolism and specific dynamic action (in the case of oxygen consumption) or postprandial nitrogen increase.
The new metabolism chamber allowed the relationship between metabolism and swimming velocity of juvenile fish with developed rheotactic response. It could be used with adult fish for similar purposes.     

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.     

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.     

水流对草鱼幼鱼趋光行为的影响

为了探讨水流对鱼类趋光性的影响, 利用自制的循环水槽装置, 以草鱼(Ctenpharyngodon idellus)幼鱼为研究对象, 研究其在光照度为300 lx, 不同流速工况(0、0.1和0.2 m/s)下的趋光性行为, 同时设黑暗静水工况为对照组。结果表明: (1) 0.2 m/s的流速可完全激发草鱼幼鱼的趋流性, 使其游泳方向多数与顶流方向呈± 20°。(2)根据草鱼幼鱼在不同流速工况下随光照度衰减在水槽内的分布情况, 计算得其在3种流速工况下的光强期望值分别为: 52.45, 34.62和37.86 lx。(3)当照度为300 lx时, 静水工况下的实验鱼在水槽中呈现“两头高, 中间低”的分布情况, 并未表现出对某一光强范围的偏好行为; 在小于感应值的低流速下, 草鱼幼鱼的分布情况总体趋势与静水工况类似, 但在远离光源处的分布较多, 多呈“逆流后退”行为; 当流速值超过感应流速时, 在趋流性的作用下, 鱼类在尾部的聚集情况明显下降, 同时在水槽中的分布更加均匀, 其原有光环境的作用减弱。研究初步证明了略大于感应值的小流速所引发的草鱼趋流性即可对其光环境响应行为产生影响。     

Temperature effects on metabolic rate, swimming performance and condition of Pacific cod Gadus macrocephalus Tilesius

Critical swimming speed ( U _crit) and rate of oxygen consumption of Pacific cod Gadus macrocephalus acclimated to 4 and 11° C were determined to assess the influence of water temperature on performance. The physiological effect of exercise trials on fish held at two temperatures was also assessed by comparing haematocrit and plasma concentrations of cortisol, metabolites and ions collected from fish before and after testing. The U _crit of fish acclimated and exercised at 4° C did not differ from those acclimated and exercised at 11° C [1·07 body lengths (total length) s⁻¹]. While the standard metabolic rate of 11° C acclimated fish was 28% higher than that of 4° C fish, no significant difference was observed between fish acclimated at the two temperatures. Plasma concentrations of cortisol, glucose and lactate increased significantly from pre- to post-swim in both groups, yet only concentrations of cortisol differed significantly between temperature treatments. Higher concentrations of cortisol in association with greater osmoregulatory disturbance in animals acclimated at the lower temperature indicate that the lower water temperature acted as an environmental stressor. Lack of significant differences in U _crit between temperature treatments, however, suggests that Pacific cod have robust physiological resilience with respect to swimming performance within temperature changes from 4 to 11° C.     

Effects of temperature on allocation of metabolic energy in perch (Perca fluviatilis) fed submaximal rations

Growth and long-term trends of oxygen consumption were monitored at 15 and 20° C in spontaneously swimming juvenile perch fed a fixed daily submaximal ration. The average rate as well as the efficiency of assimilation were the same at the two temperatures but a much higher proportion of food energy was allocated to activity and maintenance, and a correspondingly lower proportion to growth, at 20 than at 15° C. By computing average specific rates of oxygen consumption separately for all light and dark periods, it was found that at 20, but not at 15° C, the' scope for spontaneous activity' of two consecutive (dark and light) phases of a diurnal cycle was indirectly proportional to the average rate of oxygen consumption in the dark phase. This indicates that at 20° C the perch displayed partial compensation for a high metabolic rate in the dark by reducing swimming activity in the following light phase. However, the overall effect of this behaviour was probably too small to make a noticeable impact on the low conversion efficiency in the juvenile perch at 20 as compared to 15° C.