The effect of demand feeding on swimming speed and feeding responses in Atlantic salmon Salmo salar L., gilthead sea bream Sparus aurata L. and European sea bass Dicentrarchus labrax L. in sea cages

Abstract Using underwater cameras, data were collected on the feeding behaviour and swimming speeds of Atlantic salmon Salmo salar L., gilthead sea bream Sparus aurata L. and European sea bass Dicentrarchus labrax L. in sea cages. Comparisons were made between the behaviours of fish fed on demand using interactive feedback systems and those of fish fed under the standard feeding practice of each farm (control). In all three species, swimming speeds were similar before feeding , but they were significantly higher in the control regimes during feeding. When fed on demand, sea bass had reduced swimming speed just before and during feeding compared with that observed during the non‐feeding periods. Higher proportions of feeding fish were observed in the control regime cages than in fish fed on demand for all three species, indicating a greater feeding intensity during meals in the control regimes. This was further supported by observations of an increase in the density of sea bass in the upper water in the control cages during feeding. The results suggest decreased levels of competition between the on demand‐fed fish during feeding, which might be hypothesized to lead to improved growth and production efficiency in aquaculture.     

Swimming behaviour as an indicator of low growth rate and impaired welfare in Atlantic halibut (Hippoglossus hippoglossus L.) reared at three stocking densities

The Atlantic halibut (Hippoglossus hippoglossus L.) is a new species in Norwegian aquaculture. However, in ongrowing trials, halibut shows variable and generally poor rates of growth. The halibut is a flatfish that spends most of its time resting on the bottom, and it was hypothesized that high stocking densities with frequent social interactions could lead to behaviour changes, reduced appetite and impaired welfare. Halibut were kept in six tanks at low, medium, and high densities (18%, 54% and 112% bottom coverage). All fish were individually tagged with Trovan™ passive implant transponder (PIT) tags, allowing an antenna to register fish swimming at the surface. Swimming and feeding behaviour was also recorded by underwater video cameras. Individual growth rates were highly variable, but food consumption and growth rates fell significantly with increasing stocking density, while individual swimming activity rose with increasing density. Frequent “surface swimmers” had a significantly lower growth rate than fish that were seldom recorded by the PIT antennae. Surface swimming may therefore be an indicator of suboptimal growth rates and impaired welfare in reared halibut.     

From fright to anticipation: using aversive light stimuli to investigate reward conditioning in large groups of Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis>)

In this study, we demonstrate how an event that is initially frightening to Atlantic salmon is turned to a positive stimulus through habituation and associative learning. The study was carried out in four commercial sized tanks (5 m³) with near industry densities (>550 fish, 16 kg m⁻³), using a delay conditioning procedure with an aversive flashing light as the conditioned stimulus and food reward as the unconditioned stimulus. By using video image analysis of the distribution of the fish in the tanks, the changes in behaviour from trial to trial could be documented in great detail. The current study documents the change in behaviour across the individual conditioning trials, clearly showing the step-by-step nature of the transition. The salmon needed more than 26 trials to become fully habituated to the flashing light but showed clear anticipatory behaviour already after about 19 trials. This demonstrates that the learning process is a combination of habituation and associative learning.     

Providing harpacticoid copepods via floating sieve improves herring (Clupea harengus L.) larval feeding incidence

Harpacticoid copepods are being considered as alternative candidates for live feed in aquaculture, but their benthic affinity may pose problems for pelagic fish larvae. We compared the swimming behaviour and feeding incidence of herring larvae (Clupea harengus) in the presence of harpacticoid copepods (Tachidius discipes) and rotifers (Brachionus plicatilis). Additionally, we provided T. discipes via a floating sieve to improve the prey availability. The comparison was performed at 5 and 10 days post hatch (dph) via 2D‐video observations. Quantitative analyses of larval trajectories allowed the estimation of feeding behaviour through a series of indicators: swimming speed, straightness of trajectories, turning angles and swimming activities (break, sink, slow, normal, fast). The outcomes highlighted that the prey type had no significant effect on swimming speed or straightness of the swimming path. However, at 10 dph directly copepod‐fed larvae spent less time in slow but more time in the normal swimming‐state than rotifer‐fed larvae and larvae fed with Tachidius via sieve. This suggests higher energy expenditure of directly copepod‐fed larvae. Moreover, the feeding incidence was higher in larvae fed with Tachidius via sieve than directly Tachidius‐fed larvae. Thus, providing harpacticoid copepods via a floating sieve can improve the rearing of marine fish larvae.     

Optomotor response and erratic response: quantitative analysis of fish reaction to towed fishing gears

Fish were observed reacting to moving net panels in tank experiments and in trawl gears towed at sea. Two typical response behaviours of haddock, saithe, mackerel, cod and flatfish to the gear are described as optomotor response and erratic response. These two responses were analysed from TV recordings of the reacting fish and are characterised by time sequences of four parameters: swimming speed, acceleration, angular velocity and distance to the net panel. When fish display stable swimming near the net mouth as in an optomotor response, variations of swimming speed, acceleration and angular velocity are relatively low and regular in their amplitude and period. The erratic response is characterised by large variations in velocity, acceleration and angular velocity and only distance to the towed netting panel, which is positive inside and negative outside, shows progressive change. It is suggested that the fish’s process of deciding between optomotor or erratic response to the gear is based on predictable parameters that describe the stimulus like sound, light level, visibility range and object contrast, combined with the limits describing the abilities of the fish to see, hear and move. The behaviour of the observed parameters suggest that the balance between these factors in a model predicting the outcome might benefit from a form of chaos theory.     

Foraging behaviour and invasiveness: do invasive Gambusia exhibit higher feeding rates and broader diets than their noninvasive relatives?

Abstract –  Serious impacts by nonindigenous species often occur via predation. The magnitude of impact is expected to be closely tied to the invading species niche breadth. For predatory impacts, diet breadth should be particularly important. We examined the relationship between a species foraging behaviour and its invasiveness and impact by comparing the feeding behaviour of four Gambusia species, two invasive and of high impact and two noninvasive. Individual feeding rates, feeding preferences, and diet breadths were tested across three prey items in a sequence of four laboratory feeding trials. Invasive Gambusia consistently fed at higher rates, but no species differences were found in feeding preferences or diet breadth. All Gambusia preferred Daphnia , avoided Lirceus , and consumed Drosophila in proportion to their availability. Female size affected most feeding variables. Larger fish consumed more prey per unit time and were able to incorporate larger prey items into their diets, thus increasing diet breadth.     

Feeding behaviour of fast- and slow-growing strains of rainbow trout, Oncorhynchus mykiss (Walbaum), during first feeding

The development of feeding behaviour in two rainbow trout strains and their hybrid was compared at the transition from endogenous nutrition to exogenous feeding. After 60 min of acclimatization, the swimming pattern, spatial distribution, agonistic behaviour and snapping of fish were quantified by making 5-min observations on groups of five fish. Behavioural patterns were recorded in four replicate groups per treatment, before and after the introduction of dry trout feed or Artemia nauplii (actometric tests) or extracts of these (olfactometric tests). Fish of the slow growing strain were less active than those of the other two strains. Greater swimming activity observed in the fish of the fast growing strain was associated with more rapid feeding and a higher number of snapping responses. Such behavioural precocity is of interest because it could provide the basis for the growth differences between the fast and slow growing strains.     

Experimental evidence for species‐specific response to turbidity in imperilled fishes

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Effects of dissolved carbon dioxide on energy metabolism and stress responses in European seabass (Dicentrarchus labrax)

Elevated carbon dioxide concentrations reduce feed intake and growth in several fish species and induce stress responses. In this study, the effects of moderately elevated CO₂ levels on performance, energy partitioning, swimming activity and stress response in European seabass were assessed. European seabass (140.0 g) were reared under two levels of CO₂ (1.6 and 7.0 mg L^?1) and two feeding levels (FLs) (maintenance and satiation) for 60 days, and fish swimming speed was recorded. At the end of the experiment, fish were subjected to an acute stress test. Blood cortisol, glucose and lactate were analysed. Energy and nitrogen balances were quantified based on measurements of body composition and digestibility coefficients. Moderately elevated chronic CO₂ level did neither affect energy requirements for maintenance nor the utilization of digestible energy for growth. However, swimming activity data suggests that FL dependent alterations in energy partitioning took place. Blood cortisol values after the acute stress were affected by additional CO₂ exposure and this effect was also dependent on FL. The elevated CO₂ exposure of 7.0 mg L^?1 appears to act as a chronic stressor as adaptive responses took place, however, this CO₂ exposure seems to be still within the allostatic load of the fish.     

Why is cannibalism less frequent when larvae of sutchi catfish Pangasianodon hypophthalmus are reared under dim light?

Survival rates of the larvae of sutchi catfish Pangasianodon hypophthalmus are reported to be three times higher under dim conditions (0.1 lx) than those under 100 lx. In this study, larval behaviour of sutchi catfish was examined under various light intensities (<0.01, 0.1, 1, 10 and 100 lx) using a CCD camera to understand why survival rates vary under different light intensities. Five‐day‐old larvae showed significantly higher swimming activity under <0.01, 0.1 and 1 lx than those under 10 and 100 lx. On the other hand, the larvae showed significantly higher aggressive behaviour under 10 and 100 lx; swimming larvae attacked resting individuals more frequently under 10 and 100 lx than those under 0 and 0.1 lx. Aggressive behaviour was considered to induce lesions, inflicted by the sharp teeth of attacking larvae, on larval skin surfaces. It is considered that the chemical substances would generate from injured skin surfaces then acted as stimuli, causing the cannibalistic behaviour in other fish around the injured fish. This study provided evidence that the observed higher survival rates depended on lower frequency of aggressive behaviour under dark or dim conditions. It is therefore recommended that larval rearing of sutchi catfish be conducted under dim (less than 1 lx) conditions.     

Appraisal of fish behaviour based on time series of fish positions issued by a 3D array of ultrasound transducers

We present a novel low-cost analysis platform and method for monitoring the behaviour of a fish population (starry sturgeons – Acipenser stellatus) – in an experimental tank using a 3-dimensional array of ultrasound transducers. The platform provides time series of fish positions, allowing for assessment of the dynamics of geometric centre (centroid) of fish population. We demonstrate that centroid evolution represents a sensitive indicator of fish behaviour under conditions varied from normal towards non-physiological limits, e.g. pH, dissolved oxygen concentration, feeding and light.Emphasis has been given to reversible effects i.e. after normalisation of water parameters undergoing “gentle” variations, the fish behaviour returns to the normal/physiological pattern.Fish behaviour was previously assessed by several techniques (e.g. electrical impedance, video monitoring, hydro-acoustics, etc.).To our knowledge, this is the first study that uses ultrasounds to continuously monitor the three-dimensional position of fish, having also the advantage of being independent on water turbidity or light level.We stress on the capabilities of the platform and related method to assess the occurrence of abnormal fish behaviour, revealed by the change over a predefined threshold exhibited by the dynamics of the fish population exposed to stimuli, and to automatically issue a warning alert (in case of e.g. non-optimum water quality), thus making it suitable for remote, cost effective monitoring of fish behaviour supporting an optimised aquaculturing.     

不同营养状况对鲫鱼偏好游泳速度的影响

为考察不同营养状况对鲤科鱼类偏好游泳速度的影响,本研究以鲫鱼(Carassius auratus)为实验对象,(25±1.0)℃条件下设置3个不同营养状况实验组:对照组(禁食48 h)、饱食组(饱食)和饥饿组(饥饿14 d),将单尾实验鱼置于梯度流速选择仪(流速范围为11.86~65.45 cm/s,等距离划分为5个流速区域,从第一到第五流速区域流速连续增加)中拍摄1 h,采用Ethovision XT9软件分析视频资料并计算实验鱼在不同流速区域平均进入频次(F,次)、单次进入停留时间(T,s/次)和平均停留时间百分比(P_t,%)等流速选择行为指标。结果显示,对照组和饱食组的F值随水流速度上升而增加,对照组的T值不同流速区域间无差异(P0.05),而饱食组第三和第四流速区域T值均显著大于其他流速区域(P0.05),因此两实验组第三和第四流速区域的P_t均显著大于其他流速区域,其偏好游泳速度为20.12~41.30 cm/s。饥饿组流速偏好行为出现表型分化,I型实验鱼在第一流速区域P_t和T值显著大于其他流速区域(P0.05),各流速区域的F值均显著小于其他实验组(P0.05),故I型实验鱼的偏好游泳速度为11.86~15.18 cm/s。而II型实验鱼则与对照组相似,在第三和第四流速区域Pt均显著大于其他流速区域(P0.05),故II型实验鱼的偏好游泳速度为20.12~41.30 cm/s。结果表明,饱食不影响鲫鱼的偏好游泳速度,可能是饱食不影响鲫鱼的游泳能力所致,但饱食组在各流速区域间的出入频次减少而停留时间有所增加。饥饿后鲫鱼偏好游泳速度出现表型分化,I型实验鱼的偏好游泳速度降低,而II型实验鱼则无明显变化,I型和II型实验鱼偏好游泳速度的差异可能是由于在饥饿条件下二者能量节约和游泳功能维持的策略不同。     

Escape‐related behaviour of Atlantic cod,Gadus morhua L., in a simulated farm situation

Cultured fish escaping from farms represent an economic loss as well as a potential problem for wild fish populations. This study investigated the escape‐related behaviour of farmed Norwegian coastal and northeast Arctic cod (NEAC), Gadus morhua L. Six groups of fish were observed during three replicate trials in a large tank that was split into two equal parts by a net wall. The fish could move freely through an opening in the net during the trials. Three groups were not fed during the trials. The first escape occurred shortly after producing the net opening (11±10.5 min; mean±standard deviation). Norwegian coastal cod were more prone to escape than the NEAC. A starvation period of 9 days increased the number of fish on the escape side of the tank. Net biting and net inspection by the fish were frequently observed, irrespective of whether the fish were fed or not. The same fish were repeatedly found on the escape side of the tank, but the propensity for recurrent escape behaviour was not related to genotype, feeding status or size. The results suggest that other factors, such as individual variation in boldness or exploration behaviour, could affect the willingness to escape.     

Ontogenetic behaviour and swimming ability of the endangered eastern freshwater cod,Maccullochella ikei,with notes on growth and development

Abstract – Understanding the larval ecology of individual fish species is fundamental in ensuring their long‐term conservation. The endangered eastern freshwater cod, Maccullochella ikei, is endemic to the Clarence and Richmond River systems of north‐eastern New South Wales, Australia. Little is known about the behaviour of larval M. ikei in the wild, particularly before and after the swim‐up stage, and following dispersal from the nest site. The aims of this study were to quantify the swimming ability, depth selection, light preference and substrate selection of hatchling to day‐30 M. ikei under controlled laboratory conditions, and to describe its growth and development over the same period. Maccullochella ikei larvae grew constantly but not consistently during the experiment. Exogenous feeding commenced around day 12, prior to the full exhaustion of the yolk. Maximal swimming ability improved daily, but maximum swimming speed declined significantly between days 12 and 13 and remained low. Maccullochella ikei larvae were initially photonegative but were positively phototactic by day 10. Depth selection was for the benthos until day 8, beyond which time larvae dispersed to all depths when released. Substrate selection was for sand in younger larvae but changed to upstream substrates as the experiment progressed. The results of the current study suggest that the period between day 10 and day 20 is critical in the early ontogeny of M. ikei, when it switched phototrophic behaviour, transitioned from endogenous to exogenous feeding and experienced a decline in swimming ability.     

Effects of feeding conditions and size differences on aggressive behaviour and cannibalism in the Pacific bluefin tuna Thunnus orientalis (Temminck and Schlegel) larvae

We examined the effects of feeding conditions and size differences on the aggressive behaviour and cannibalism in Pacific bluefin tuna (PBT) Thunnus orientalis larvae. In a 24 h experiment, restricted feeding alone was found to remarkably increase the frequency of aggressive behaviour, which was further elevated by differences in fish size. In a 4‐day rearing experiment, while aggressive behaviour was increased by restricted feeding alone, the frequency of cannibalism did not change significantly. Although the frequency of aggressive behaviour did not increase with difference in size factor, small fish in this group gradually tended to die over 4‐day period. In the restricted feeding and size difference group, large fish were observed to bite the small fish, and almost all the small fish died on the day after the start of the experiment. These results suggest that the aggressive behaviour of PBT larvae is chiefly increased by the shortage of live food; however, deaths related to cannibalism mainly occurred in small larvae and rapidly increased with food restriction and differences in fish size.     

Submerged light increases swimming depth and reduces fish density of Atlantic salmon Salmo salar L. in production cages

Artificial photoperiods that postpone sexual maturation and increase growth are now widely used in the Atlantic salmon Salmo salar L. farming industry. Few studies have been carried out to examine the effect of this treatment on fish behaviour and welfare in production cages. In this study, echo‐integration was used to observe the swimming depth and fish density of salmon in 20‐m‐deep production cages illuminated by lamps mounted above the water surface (SURF) or submerged in the cage (SUBS). From January to May, SUBS swam at a greater depth (5–11 m) than SURF (1–3 m) at night. SURF descended and SUBS ascended at dawn, but SUBS were still swimming at greater depth than SURF during the day from January to March. The difference in swimming depth resulted in SURF swimming at a median fish density about twice as high as SUBS at night and up to five times the calculated fish density. SURF increased the utilization of the cage volume as the biomass increased, but fish swimming at the highest density did so at up to 20 times the calculated fish density. The results suggest that salmon position themselves in relation to the artificial light gradient to maintain schooling behaviour and that the use of submersible lights may be a precaution to secure the welfare of caged salmon.     

Monitoring feeding behaviour and food intake: methods and applications

There are three main methods used in the study of feeding behaviour and food consumption in captive populations of fish. These are direct observation, the recording of feeding activity using‘on-demand’feeders, and the quantitative determination of gastrointestinal content using a non-invasive‘labelled feed’technique. Observational techniques have been widely used in studies of foraging behaviour and feeding responses, but these methods are usually confined to the study of fish held in small groups.‘Labelled feed’techniques are well suited to the monitoring of feed intake of individual fish held in large groups, but cannot be used for the continuous monitoring of feeding behaviour. They are, therefore, not particularly suitable for the collection of information about short-term changes in feeding rhythms, or possible diel changes in feeding activity. Studies of feeding activity may best be carried out using computer-operated‘on-demand’feeding systems, but the data collected using these systems give no information about individual feed consumption. Examples are presented showing how data collected using the three methods can be used to provide insights into how various abiotic and biotic factors influence fish feeding.     

Influence of feeding regime on intraspecific competition, fin damage and growth in 1+ Atlantic salmon parr (Salmo salar L.) held in freshwater production cages

The feeding behaviour, growth and feed conversion ratio (FCR) of cage‐held Atlantic salmon parr (Salmo salar L.) were studied when in 576 m³ (12 m × 12 m × 4 m) commercial freshwater cages under ambient water temperature (8.84±3.53°C) and photoperiod (11.02±2.05 h) for 205 days. The effect of feeding regime on fin damage was also investigated. Six groups (n=31 234±2051 fish group⁻¹, initial stocking density 1.25±0.14 kg m⁻³) were fed to satiation using either (a) an imposed regime involving scheduled, fixed ration feeding every 10 min from dawn till dusk or (b) on demand from dawn till dusk using commercial interactive feedback systems. During feeding, there were no significant differences in aggression although swimming speeds and turning angles were significantly higher in fish under the imposed regime. On‐demand feeding significantly reduced the incidence of dorsal fin damage. There was no clear relationship between fish size, feed regime and the incidence of fin damage until 1 week before the fish were transferred to marine cages, when the smallest fish under each feeding regime had the highest incidence of fin damage. Interestingly, growth did not differ between regimes, but fish under the imposed regime were significantly overfed and achieved higher FCRs.     

School for learning: sharing and transmission of feeding information in jack mackerel <Emphasis Type="Italic">Trachurus japonicus</Emphasis> juveniles

We aimed to evaluate the effect of social interaction on learning in juvenile jack mackerel Trachurus japonicus. We first compared the learning process between single fish and groups of fish. Reward-conditioned learning was established in eight trials in both treatments, whereas individuals in the group treatment responded to stimuli more frequently in the feeding area than in the single fish. This implies that information about the feeding area was shared in the group and pursuing other individuals gave them a behavioral advantage for feeding. We then investigated whether information on the feeding area can be transmitted through observation of other individuals in aligned tanks. Fish in the control group required six trials to be conditioned to aeration stimuli and feeding location, whereas those in the observation treatment required only three trials for this learning. This result implies that information on the feeding area was transmitted through visual observations. The present research suggests that sharing and transmission of information occur in schools of jack mackerel. Schooling behavior would thus enable optimization of the foraging behavior in this species.     

Consumption of feeds containing the antibiotic sulfadiazine by gilthead sea bream (Sparus aurata,L.) and rainbow trout (Onchorynchus mykiss,Walbaum)

A method for studying the efficacy of sulfadiazine as a marker to estimate feed intake is presented. Feeding studies were carried out with two species, rainbow trout as a freshwater fish model and gilthead sea bream as a marine fish model, using two temperatures and two sizes of juveniles. The study showed the different feeding behaviour observed in both species, depending on the temperature and the number of days feeding the particles with sulfadiazine, and confirmed a dominant‐subordinate behaviour especially in the case of gilthead sea bream juveniles.