Less is more: density influences the development of behavioural life skills in trout

Theory suggests that habitat structure and population density profoundly influence the phenotypic development of animals. Here, we predicted that reduced rearing density and increased structural complexity promote food search ability, anti-predator response and the ability to forage on novel prey, all behavioural skills important for surviving in the wild. Brown trout were reared at three densities (conventional hatchery density, a fourth of conventional hatchery density and natural density) in tanks with or without structure. Treatment effects on behaviour were studied on trout fry and parr, whereupon 20 trout from each of the six treatment groups were released in an enclosed natural stream and recaptured after 36 days. Fry reared at natural density were faster to find prey in a maze. Moreover, parr reared at natural density were faster to eat novel prey, and showed more efficient anti-predator behaviour than fish reared at higher densities. Furthermore, parr reared at reduced densities were twice as likely to survive in the stream as trout reared at high density. In contrast, we found no clear treatment effects of structure. These novel results suggest that reduced rearing densities can facilitate the development of behavioural life skills in captive animals, thereby increasing their contribution to natural production.     

Evaluation of a new semi-natural incubation technique for Atlantic salmon Salmo salar fry

The incubation environment had a significant effect on fork length ( L _F) and body mass ( M ) of Atlantic salmon Salmo salar fry at the time of emergence. Fry that were incubated in a new incubator design, that mimics the conditions in a natural redd (the Bamberger-Box), achieved significantly greater and attained a significantly higher M than those reared in conventional hatchery troughs for control. Fewer fry from the Bamberger-Boxes had visible deformities compared with those from the hatchery troughs. Results were consistent for five consecutive seasons using both wild and domesticated broodstock from genetically different origins. Survival from the eyed embryo stage in the Bamberger-Boxes and hatchery troughs was >93% during normal climatic conditions. Only larvae reared in Bamberger-Boxes, however, survived abnormally high water temperatures during one test season. The results demonstrate that the Bamberger-Box is an effective alternative to the conventional incubation technology.     

Environmental rearing conditions produce forebrain differences in wild Chinook salmon Oncorhynchus tshawytscha

Recent studies suggest that hatchery-reared fish can have smaller brain-to-body size ratios than wild fish. It is unclear, however, whether these differences are due to artificial selection or instead reflect differences in rearing environment during development. Here we explore how rearing conditions influence the development of two forebrain structures, the olfactory bulb and the telencephalon, in juvenile Chinook salmon (Oncorhynchus tshawytscha) spawned from wild-caught adults. First, we compared the sizes of the olfactory bulb and telencephalon between salmon reared in a wild stream vs. a conventional hatchery. We next compared the sizes of forebrain structures between fish reared in an enriched NATURES hatchery and fish reared in a conventional hatchery. All fish were size-matched and from the same genetic cohort. We found that olfactory bulb and telencephalon volumes relative to body size were significantly larger in wild fish compared to hatchery-reared fish. However, we found no differences between fish reared in enriched and conventional hatchery treatments. Our results suggest that significant differences in the volume of the olfactory bulb and telencephalon between hatchery and wild-reared fish can occur within a single generation.     

Impact of live food on survival and growth of hatchery‐reared sea trout (Salmo trutta trutta L.) parr in the wild

Survival rates and growth parameters of hatchery‐reared sea trout (Salmo trutta trutta L.) fry were determined after stocking in the wild. The larvae were hatchery‐reared for 12 weeks in two groups: fry were fed either on live zooplankton and live chironomidae larvae (LFG), or fed a pellet diet (PFG). The survival rate and specific growth rates were higher in the LFG than in the PFG group. Most effective for hatchery‐reared fish intended for stocking was the natural, live feed. The mean number of chironomid larvae found in the stomachs of fish that were initially captured in the wild was significantly higher in the LFG than in the PFG group. The live diet supplied in the rearing period had a positive impact on the foraging skills of the sea trout fry and their survival in the wild after their release on 24 April 2010.     

Can structural enrichment reduce predation mortality and increase recaptures of hatchery-reared Atlantic salmon Salmo salar L. fry released into the wild?

Captive-reared fish often have poor survival in the wild and may fail to boost threatened populations. Enrichment during the nursery period can in some circumstances generate a broader behavioural repertoire than conventional hatchery production. Yet, we do not know if enrichment promotes survival after release into the wild. We conducted a field experiment during three field seasons using age 0+ year Atlantic salmon Salmo salar to investigate if enrichment during rearing, in the form of structural complexity (shelters), reduced immediate (within 2 days after release) predation mortality by piscine predators (brown trout Salmo trutta) and if such rearing environments improved long-term (2–3 months after release) post-release survival. In addition, we investigated if predation mortality of released fry was size-selective. S. salar fry were reared in a structurally enriched environment or in a conventional rearing environment and given otolith marks using alizarin during the egg stage to distinguish between enriched and conventionally-reared fry. The outcome from the field experiments showed that structural enrichment did not consistently reduce immediate predation mortality and it did not improve, or had a negative effect on, the recapture rate of fry from the river 2–3 months after release. The data also showed that enriched rearing tended to reduce growth. Additionally, we found that S. trutta predators fed on small individuals of the released fry. Overall, the data suggest that structural enrichment alone is not sufficient to improve long-term survival of hatchery-reared fish after release and that other factors might affect post-release survival.     

Effects of habitat enrichment and food availability on the foraging behaviour of juvenile Atlantic Cod (<Emphasis Type="Italic">Gadus morhua L</Emphasis>)

The environment can play an important role in shaping how an animal behaves, and how well the animal performs in a particular environment can be influenced by early experiences. The tradition of releasing captive-reared juveniles into the wild in an effort to strengthen wild fish populations has often had little success owing to high post-release mortality. Fish reared under standard hatchery conditions are provided with fewer stimuli and they receive excess quantities of pellet food that are easy to handle and consume. Captive reared fish therefore appear to be under-stimulated and overfed. Several studies have demonstrated that simple structural enrichment in the rearing facilities promotes flexible behaviour compared to fish reared in plain, standard hatchery tanks. Less attention has been given to the effects of the diet. Here we use a cross-factored design to test the relative role of food ration and spatial enrichment on foraging behaviour. Our results show that fish from enriched environments, regardless of previous food-ration size, were more reluctant to start feeding on the first day in a novel arena. On day two and three, however, fish with prior experience of a low food ration showed greater foraging activity and efficiency than fish fed on full rations. On the second and third day, prior experience with enrichment was less important. We discuss how early feeding experience in combination with structural enrichment may contribute in producing fish that are better suited for release into the wild.     

Silver spoons in the rough: can environmental enrichment improve survival of hatchery Atlantic salmon Salmo salar in the wild?

This study tested the ‘silver spoon’ hypothesis which posits that individuals that develop under favourable conditions should enjoy a fitness advantage later in life because they are more likely to recognize and settle in high‐quality habitats. Atlantic salmon Salmo salar of two age classes (0+ and 1+ years) were reared in environmentally enriched or standard hatchery tanks for a short period (c. 10 weeks), were then released into a natural river and sampled on repeated occasions to test for silver‐spoon effects. Compared with controls, enriched fish had a 6·4% higher recapture rate and settled in higher velocity habitats when they were stocked as 0+ year fry, but not when they were stocked as 1+ year parr. The opportunity for selection was generally higher for environmentally enriched fish than for controls, and also higher for 0+ than for 1+ year fish. Selection favoured individuals with high condition factor, extensive fat reserves and longer than average pectoral fins in both age classes but favoured a small body size in 1+ year and a large body size in 0+ year releases. Stomach analysis showed that enriched fish ate more, and adapted quicker to natural prey than controls. These results provide support for silver‐spoon effects in fish and indicate that enrichment can improve post‐release performance in conservation programmes, but seemingly only if fish are not kept in captivity for too long.     

Inside the heads of David and Goliath: environmental effects on brain morphology among wild and growth-enhanced coho salmon Oncorhynchus kisutch

Transgenic and wild-type individual coho salmon Oncorhynchus kisutch were reared in hatchery and near-natural stream conditions and their brain and structure sizes were determined. Animals reared in the hatchery grew larger and developed larger brains, both absolutely and when controlling for body size. In both environments, transgenics developed relatively smaller brains than wild types. Further, the volume of the optic tectum of both genotypes was larger in the hatchery animals and the cerebellum of transgenics was smaller when reared in near-natural streams. Finally, wild types developed a markedly smaller telencephalon under hatchery conditions. It is concluded that, apart from the environment, genetic factors that modulate somatic growth rate also have a strong influence on brain size and structure.     

Stocking experiment with Atlantic salmon and sea trout parr reared on either live prey or a pellet diet

Survival rate and growth parameters of Atlantic salmon fry and sea trout fry were determined after stocking in the wild. Before release (22 May 2009) into the wild the larvae were reared for 10 weeks in the hatchery in three groups: (i) fry fed on live zooplankton , (ii) fry fed on larvae of live nekton, and (iii) fry fed on prepared pellet food. In autumn (15 September 2010) the fish were caught in the wild; the survival rate and growth parameters of both Atlantic salmon and sea trout were the highest in the zooplankton‐fed group, whilst the pellet‐fed group had the lowest survival rate and growth value parameters. Most effective food for hatchery‐reared fishes to be used as stock was the natural living zooplankton. The general conclusion is that the live diet supplied in the rearing period has a positively impact on fish survival in the wild.     

The food web positioning and trophic niche of the non-indigenous round goby: a comparison between two Baltic Sea populations

Native species may show invasiveness toward a recipient ecosystem through increases in abundance as a result of artificial stocking events. Salmonid species are typical examples of native invaders whose abundance is increased after stocking with hatchery fish. This study evaluated the effects of hatchery chum salmon fry on sympatric wild masu salmon fry, benthic invertebrate prey, and algae, after a single stocking event in Mamachi stream, Hokkaido, northern Japan. The results suggested that the stocked hatchery chum salmon fry decreased the foraging efficiency and growth of the wild masu salmon fry through interspecific competition, and depressed the abundance of Ephemerellidae and total grazer invertebrates (Glossosomatidae, Heptageniidae, and Baetidae) through predation. Also, the hatchery chum salmon fry may increase algal biomass through depression of grazer abundance by predation (top-down effect). These results suggested that a single release of hatchery chum salmon fry into a stream may influence the recipient stream ecosystem.     

Predator‐induced hyperventilation in wild and hatchery Atlantic salmon fry

Following exposure to a predator stimulus (a brown trout Salmo trutta ), the opercular rate of Atlantic salmon Salmo salar fry increased by 35·3 ± 11·0%(mean ± 95% CI). The time taken for opercular rate to decline to baseline levels depended upon the occurrence of any associated locomotory activity. Opercular rates of fish that dashed when exposed remained elevated for 38·2 ± 20·6 min, whereas those of individuals that did not move ('freezers') recovered within 7·2 ± 2·9 min. The duration that opercular rate remained elevated was positively correlated with the magnitude of the elevation, which was higher in 'dashers' than freezers. The maximum opercular rate in 'freezers' was similar between wild fry and hatchery‐reared fry (from wild parents). There was a significant delay, however, in hatchery compared with wild fry in the time until peak ventilatory response and onset in the decline phase. This difference in opercular response suggests that hatchery fish were slower to realize fully the potential danger from the predator. Any delay in response could be directly attributed to the effect of hatchery‐rearing environment, rather than domestication or hatchery selection effects.     

Rearing environment affects spatial learning in juvenile Chinook salmon Oncorhynchus tshawytscha

We tested the prediction that a complex physical rearing environment would enhance short-term spatial memory as assessed by learning ability in a spatial navigation task in juvenile Chinook salmon Oncorhynchus tshawytscha. We reared fish in two low-density treatments, where fish were either in bare fiberglass tanks (bare) or in tanks with physical structure (complex). We also tested conventionally reared high-density hatchery fish to compare with these other experimental treatments. Our reason for including this third hatchery treatment is that the two low-density treatments, aside from the manipulation of structure, followed a rearing programme that is designed to produce fish with more wild-like characteristics. We tested individually marked fish for seven consecutive days and recorded movement and time to exit a testing maze. Stimulus conspecific fish outside the exit of the maze provided positive reinforcement for test fish. Fish from the bare treatment were less likely to exit the start box compared with fish in the complex and hatchery treatments. However, fish in the hatchery treatment were significantly more likely to exit the maze on their own compared with both the bare and complex treatments. Hatchery fish effectively learned the task as shown by a decrease in the number of mistakes over time, but the number of mistakes was significantly greater on the first day of trials. Increasing habitat complexity with structure may not necessarily promote spatial learning ability, but differences between hatchery and experimental treatments in rearing density and motivation to be near conspecifics likely led to observed behavioural differences.     

Effects of environmental enrichment on survivorship,growth, sex ratio and behaviour in laboratory maintained zebrafish Danio rerio

Environmental enrichment involves increasing the complexity of a fish's environment in order to improve welfare. Researchers are legally obliged to consider the welfare of laboratory animals and poor welfare may result in less robust data in experimental science. Laboratory zebrafish Danio rerio are usually kept in bare aquaria for ease of husbandry and, despite being a well-studied species, little is known about how laboratory housing affects their welfare. This study shows that environmental enrichment, in the form of the addition of gravel substratum and plants into the tank, affects survivorship, growth and behaviour in laboratory-maintained D. rerio. Larvae reared in enriched tanks had significantly higher survivorship compared with larvae reared in bare tanks. Effects of the tank conditions on growth were more variable. Females from enriched tanks had a higher body condition than females maintained in bare tanks, but intriguingly this was not the case for males, where the only difference was a more variable body condition in males maintained in bare tanks. Sex ratio in the rearing tanks did not differ between treatments. Resource monopolisation was higher for fish in enriched tanks than for those in bare tanks. Fish from enriched tanks displayed lower levels of behaviours associated with anxiety compared with fish from bare tanks when placed into a novel environment. Thus, this study demonstrates differences in welfare for D. rerio maintained under different environmental conditions with enhancements in welfare more commonly associated with tank enrichment.     

Genetic versus Rearing-Environment Effects on Phenotype: Hatchery and Natural Rearing Effects on Hatchery- and Wild-Born Coho Salmon

With the current trends in climate and fisheries, well-designed mitigative strategies for conserving fish stocks may become increasingly necessary. The poor post-release survival of hatchery-reared Pacific salmon indicates that salmon enhancement programs require assessment. The objective of this study was to determine the relative roles that genotype and rearing environment play in the phenotypic expression of young salmon, including their survival, growth, physiology, swimming endurance, predator avoidance and migratory behaviour. Wild- and hatchery-born coho salmon adults (Oncorhynchus kisutch) returning to the Chehalis River in British Columbia, Canada, were crossed to create pure hatchery, pure wild, and hybrid offspring. A proportion of the progeny from each cross was reared in a traditional hatchery environment, whereas the remaining fry were reared naturally in a contained side channel. The resulting phenotypic differences between replicates, between rearing environments, and between cross types were compared. While there were few phenotypic differences noted between genetic groups reared in the same habitat, rearing environment played a significant role in smolt size, survival, swimming endurance, predator avoidance and migratory behaviour. The lack of any observed genetic differences between wild- and hatchery-born salmon may be due to the long-term mixing of these genotypes from hatchery introgression into wild populations, or conversely, due to strong selection in nature—capable of maintaining highly fit genotypes whether or not fish have experienced part of their life history under cultured conditions.     

Environmental effects on behavioural development consequences for fitness of captive‐reared fishes in the wild

Why do captive‐reared fishes generally have lower fitness in natural environments than wild conspecifics, even when the hatchery fishes are derived from wild parents from the local population? A thorough understanding of this question is the key to design artificial rearing environments that optimize post‐release performance, as well as to recognize the limitations of what can be achieved by modifying hatchery rearing methods. Fishes are generally very plastic in their development and through gene–environment interactions, epigenetic and maternal effects their phenotypes will develop differently depending on their rearing environment. This suggests that there is scope for modifying conventional rearing environments to better prepare fishes for release into the wild. The complexity of the natural environment is impossible to mimic in full‐scale rearing facilities. So, in reality, the challenge is to identify key modifications of the artificial rearing environment that are practically and economically feasible and that efficiently promote development towards a more wild‐like phenotype. Do such key modifications really exist? Here, attempts to use physical enrichment and density reduction to improve the performance of hatchery fishes are discussed and evaluated. These manipulations show potential to increase the fitness of hatchery fishes released into natural environments, but the success is strongly dependent on adequately adapting methods to species and life stage‐specific conditions.     

Do stocked hatchery‐reared juveniles ecologically suppress wild juveniles in Salvelinus leucomaenis?

The dominancy of semi‐wild and hatchery‐reared white‐spotted charr Salvelinus leucomaenis juveniles was evaluated using pair‐wise enclosure tests and field stocking tests. The semi‐wild S. leucomaenis originated in a hatchery, being stocked into the test stream as eyed‐eggs. In the pair‐wise enclosure test, the semi‐wild S. leucomaenis dominated the hatchery S. leucomaenis that were of a similar standard length (L_S). The semi‐wild S. leucomaenis were subordinate to hatchery S. leucomaenis that were > 11% larger in L_S. In the field stocking test, the abundance and growth of semi‐wild S. leucomaenis was decreased in the presence of larger hatchery S. leucomaenis (14% larger L_S). Taken together, these results suggest that larger hatchery S. leucomaenis ecologically suppress the smaller semi‐wild S. leucomaenis. Salvelinus leucomaenis juveniles that are stocked with the intention of supplementing natural populations should be < 10% larger than their wild counterparts at the time of stocking to minimize their competitive advantage. The semi‐wild and hatchery S. leucomaenis used in both tests were genetically similar individuals, suggesting that the differences are due to the early rearing environment of either a natural stream or hatchery. The hatchery S. leucomaenis have lower levels of aggression as a result of selection in the hatchery rearing environment. Rearing in a natural stream from the eyed‐egg stage is likely to increase their lowered aggression.     

Environmental plasticity and colonisation history in the Atlantic salmon microbiome: A translocation experiment

Microbial communities associated with the gut and the skin are strongly influenced by environmental factors, and can rapidly adapt to change. Historical processes may also affect the microbiome. In particular, variation in microbial colonisation in early life has the potential to induce lasting effects on microbial assemblages. However, little is known about the relative extent of microbiome plasticity or the importance of historical colonisation effects following environmental change, especially for nonmammalian species. To investigate this we performed a reciprocal translocation of Atlantic salmon between artificial and semi‐natural conditions. Wild and hatchery‐reared fry were transferred to three common garden experimental environments for 6 weeks: standard hatchery conditions, hatchery conditions with an enriched diet, and simulated wild conditions. We characterized the faecal and skin microbiome of individual fish before and after the environmental translocation, using a BACI (before‐after‐control‐impact) design. We found evidence of extensive microbiome plasticity for both the gut and skin, with the greatest changes in alpha and beta diversity associated with the largest changes in environment and diet. Microbiome richness and diversity were entirely determined by environment, with no detectable effects of fish origin, and there was also a near‐complete turnover in microbiome structure. However, we also identified, for the first time in fish, evidence of historical colonisation effects reflecting early‐life experience, including ASVs characteristic of captive rearing. These results have important implications for host adaptation to local selective pressures, and highlight how conditions experienced during early life can have a long‐term influence on the microbiome and, potentially, host health.     

Role of diet and experience in the development of feeding behaviour in largemouth bass, Micropterus salmoides

Two groups of largemouth bass, Micropterus salmoides , were reared in the laboratory. One group was reared on an artificial, passive diet (frozen brine shrimp) whereas the second was reared on a natural, active diet (cultured zooplankton). Observations on the development of feeding behaviour indicated that the motor patterns and duration (number of weeks in the behavioural repertoire) of the feeding acts did not differ between fry reared on the two diets. While feeding on their respective diets, natural-diet fry performed significantly more orientations and bites, the two major early feeding acts, than did the artificial-diet fry. When tested with live fish prey, fish reared on the natural diet performed fewer orientations and strikes and captured more prey per fry than did the artificial-diet fry. Natural-diet fry had a significantly better net efficiency (captures minus strikes minus orientations) than did artificial-diet fry. Diet, experience, and length (T.L.) of fry affected their predator efficiency significantly. We argue that providing hatchery-reared bass fry with an opportunity to prey on live forage fish once or twice before their release would enhance their survival and eventual recruitment into natural populations.     

Egg, fry and smolt production from salmon, Salmo salar L. and sea trout Salmo trutta L. reared entirely in fresh water

For five years hatchery reared salmon and sea trout which had spent their entire lives in hatchery tanks were stripped each Autumn to investigate the survival of the resulting offspring.