Effects of Hatchery Rearing on Brain Structures of Rainbow Trout, Oncorhynchus mykiss

In this study, we contrast brain morphology from hatchery and wild reared stocks to examine the hypothesis that in salmonid fishes, captive rearing produces changes in brain development. Using rainbow trout, Oncorhynchus mykiss, as a model, we measured eight regions of the salmonid brain to examine differences between wild and hatchery reared fish. We find using multiple analysis of covariance (MANCOVA), analysis of covariance (ANCOVA) and discriminant function analysis (DFA) that the brains of hatchery reared fish are relatively smaller in several critical measures than their wild counterparts. Our work may suggest a mechanistic basis for the observed vulnerability of hatchery fish to predation and their general low survival upon release into the wild. Our results are the first to highlight the effects of hatchery rearing on changes in brain development inbreak fishes.     

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.     

Evidence for Morphometric Differentiation of Wild and Captively Reared Adult Coho Salmon: A Geometric Analysis

As part of a comprehensive genetic evaluation of reproduction in naturally spawning coho salmon, Oncorhynchus kisutch, we examined morphometric variation in captively reared and wild adults from Hood Canal, Washington (U.S.A.) for evidence of differentiation between these groups. We collected captively reared fish as parr from two stocks and reared to adulthood at a freshwater hatchery, maturing in 1995 and 1996; we sampled closely size-matched wild fish as they returned to a neighboring stream in both years. Multivariate analysis of shape variation by Procrustes coordinates, visualized by thin-plate splines, indicated that the captively reared adults were differentiated from the wild fish by sharply reduced sexual dimorphism as well as smaller heads and less hooked snouts, increased trunk depth, larger caudal peduncles, shorter dorsal fins, larger hindbodies and a reduction in body streamlining. The differences between the captively reared and wild fish were similar to but more pronounced than some differences previously reported between hatchery and wild coho salmon. The magnitude and pattern of differences suggested that at least some of them were environmentally induced. Shape variation showed an allometric relationship with variation in body (measured as centroid) size. Morphometric variation was a poor correlate of most spawning behaviors. Nevertheless, our results suggest that the morphometric consequences of captive rearing for mate selection and reproductive activity of spawning fish may limit its effectiveness as a restorative tool.     

Rearing Environment Affects the Brain Size of Guppies: Lab-Reared Guppies have Smaller Brains than Wild-Caught Guppies

Animals bred for captivity often have smaller brains and behave differently than their wild counterparts. These differences in brain size have been attributed to genetic changes resulting from, for example, inbreeding depression and pleiotropic effects of artificial selection for traits such as docility. A critical question, though, is whether these differences in brain size are due to plastic responses to the environment, not just genetic changes. We observed a large reduction in brain size in first generation, lab-reared female guppies compared with wild-caught ones (19% smaller telencephalon, 17% smaller optic tectum). We then reared first-generation, lab-born guppies in environments varying in spatial complexity and size in an attempt to isolate factors that might increase brain size and change temperament, but no significant differences in phenotype were observed. The results of these experiments show that, although the environmental factors responsible for the effect have not been found, even first generation lab-reared individuals can have smaller brains than wild individuals.     

Habitat-dependent and -independent plastic responses to social environment in the nine-spined stickleback (Pungitius pungitius) brain

The influence of environmental complexity on brain development has been demonstrated in a number of taxa, but the potential influence of social environment on neural architecture remains largely unexplored. We investigated experimentally the influence of social environment on the development of different brain parts in geographically and genetically isolated and ecologically divergent populations of nine-spined sticklebacks (Pungitius pungitius). Fish from two marine and two pond populations were reared in the laboratory from eggs to adulthood either individually or in groups. Group-reared pond fish developed relatively smaller brains than those reared individually, but no such difference was found in marine fish. Group-reared fish from both pond and marine populations developed larger tecta optica and smaller bulbi olfactorii than individually reared fish. The fact that the social environment effect on brain size differed between marine and pond origin fish is in agreement with the previous research, showing that pond fish pay a high developmental cost from grouping while marine fish do not. Our results demonstrate that social environment has strong effects on the development of the stickleback brain, and on the brain''s sensory neural centres in particular. The potential adaptive significance of the observed brain-size plasticity is discussed.     

Exposure to predators does not lead to the evolution of larger brains in experimental populations of threespine stickleback

Natural selection is often invoked to explain differences in brain size among vertebrates. However, the particular agents of selection that shape brain size variation remain obscure. Recent studies suggest that predators may select for larger brains because increased cognitive and sensory abilities allow prey to better elude predators. Yet, there is little direct evidence that exposure to predators causes the evolution of larger brains in prey species. We experimentally tested this prediction by exposing families of 1000–2000 F2 hybrid benthic‐limnetic threespine stickleback to predators under naturalistic conditions, along with matched controls. After two generations of selection, we found that fish from the predator addition treatment had significantly smaller brains (specifically smaller telencephalons and optic lobes) than fish from the control treatment. After an additional generation of selection, we reared experimental fish in a common environment and found that this difference in brain size was maintained in the offspring of fish from the predator addition treatment. Our results provide direct experimental evidence that (a) predators can indeed drive the evolution of brain size–‐but not in the fashion commonly expected and (b) that the tools of experimental evolution can be used to the study the evolution of the vertebrate brain.     

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.     

Genetic implications of hatchery rearing in Atlantic salmon: effects of rearing environment on genetic composition

In an experiment to investigate genetic consequences of hatchery rearing in salmon, allozyme variation at five polymorphic loci was examined in Atlantic salmon of known initial genetic composition, which were reared throughout freshwater life in the hatchery or stocked into the wild as swim-up fry. The genetic composition of the juveniles in the hatchery remained homogeneous from fertilization up to stocking, and from stocking to 2+ in the wild, however, those remaining at the hatchery developed genetic differences among smolting and nonsmolting 1+ parr. These differences were attributed to conditions leading to early smolting at 1+ among the hatchery fish, with 1+ smolts diverging from the gene pool from which they were derived, whereas those stocked into the wild did not smolt until a year later and retained the original genetic composition. The results are discussed in relation to hatchery rearing of salmon and implications for the use of reared fish in stocking and enhancement programmes.     

Evidence for small scale variation in the vertebrate brain: mating strategy and sex affect brain size and structure in wild brown trout (Salmo trutta)

The basis for our knowledge of brain evolution in vertebrates rests heavily on empirical evidence from comparative studies at the species level. However, little is still known about the natural levels of variation and the evolutionary causes of differences in brain size and brain structure within‐species, even though selection at this level is an important initial generator of macroevolutionary patterns across species. Here, we examine how early life‐history decisions and sex are related to brain size and brain structure in wild populations using the existing natural variation in mating strategies among wild brown trout (Salmo trutta). By comparing the brains of precocious fish that remain in the river and sexually mature at a small size with those of migratory fish that migrate to the sea and sexually mature at a much larger size, we show, for the first time in any vertebrate, strong differences in relative brain size and brain structure across mating strategies. Precocious fish have larger brain size (when controlling for body size) but migratory fish have a larger cerebellum, the structure in charge of motor coordination. Moreover, we demonstrate sex‐specific differences in brain structure as female precocious fish have a larger brain than male precocious fish while males of both strategies have a larger telencephalon, the cognitive control centre, than females. The differences in brain size and structure across mating strategies and sexes thus suggest the possibility for fine scale adaptive evolution of the vertebrate brain in relation to different life histories.     

Population characteristics of pallid sturgeon (Scaphirhynchus albus (Forbes & Richardson, )) in the Lower Missouri River

Population characteristics of pallid sturgeon Scaphirhynchus albus in the lower Missouri River are relatively unknown. Therefore, data collected from the Nebraska Game and Parks Commission Pallid Sturgeon Population Assessment Program was synthesized to (i) document the population structure of pallid sturgeon by origin (hatchery‐reared or wild), gender, and reproductive readiness, (ii) document the minimum size and age‐at‐maturity by gender, and (iii) document the fecundity rates of the fish that were successfully spawned in the hatchery. During this 4‐year study (2008–2011), relative abundance for wild and hatchery‐reared pallid sturgeon collected with gill nets did not significantly change whereas relative abundance for wild fish using trot lines declined significantly. The proportion of hatchery‐reared pallid sturgeon increased annually, with the population being composed primarily of hatchery‐reared fish. The proportion of reproductively ready females to non‐reproductively ready females was 1 : 2.0, compared to male ratios at 1 : 0.9. Minimum fork length‐at‐maturity was estimated for females at 788 mm and for males at 798 mm. Minimum age‐at‐maturity for hatchery‐reared released fish was age‐9 for females and age‐7 for males. Highest relative fecundity, based on the ovosomatic index, was 10% with an overall mean of 7%. The number of eggs per ml (egg size) was not correlated with fork length (P = 0.0615) or weight (P = 0.0957). Relative condition factor (Kn) for females was significantly different by reproductive condition (P = 0.0014) and Kn for males did not differ between reproductive conditions (P = 0.2634). Detecting shifts in population characteristics are essential not only to understand population dynamics since hatchery inputs and natural perturbations continue to change the population structure but also to assess species recovery efforts to ensure long‐term species sustainability.     

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.     

Rearing and migration of juvenile Chinook salmon (<Emphasis Type="Italic">Oncorhynchus tshawytscha</Emphasis>) in a large river floodplain

Off-channel habitat has become increasingly recognized as key for migratory fishes such as juvenile Chinook salmon (Oncorhynchus tshawytscha). Hence, floodplain habitat has been identified as critical for the continued persistence of California’s Central Valley salmon, particularly the Yolo Bypass, the primary floodplain of the Sacramento River. To provide insight into factors supporting juvenile salmon use of this 240 km², partially leveed floodplain, we examined inter- and intra-annual relationships between environmental correlates and residency time, apparent growth, emigration, migratory phenotype, and survival over more than a decade for natural-origin (“wild”) fish and experimentally-released hatchery fish. Flood duration was positively associated with hatchery juveniles residing longer and achieving larger size. Wild juveniles grew larger and emigrated later with cumulative temperature experience (accumulated thermal units) and warmer average annual temperatures during flood years. Within years, both wild and hatchery salmon departed the floodplain as flood waters receded. Parr-sized juveniles dominated outmigrant composition, though fry and smolt-sized juveniles were also consistently observed. Survival to the ocean fishery was not significantly different between hatchery fish that reared in the Yolo Bypass versus those that reared in the main stem Sacramento River. Our study indicates improved frequency and duration of connectivity between the Sacramento River and the Yolo Bypass could increase off-channel rearing opportunities that expand the life history diversity portfolio for Central Valley Chinook salmon.     

Are antipredator behaviours of hatchery Salmo salar juveniles similar to wild juveniles?

This study explores how antipredator behaviour of juvenile Atlantic salmon Salmo salar developed during conventional hatchery rearing of eggs from wild brood stock, compared with the behaviour of wild‐caught juveniles from the same population. Juveniles aged 1+ years were tested in two unfamiliar environments; in one S. salar were presented with simulated predator attacks and in the other they were given the opportunity to explore an open‐field arena. No difference was found in their spontaneous escape responses or ventilation rate (reflex responses) after simulated predator attacks. Hatchery‐reared juveniles were more risk‐prone in their behaviours than wild‐caught individuals. Hatchery juveniles stayed less time in association with shelter. In the open‐field arena, hatchery juveniles were more active than wild juveniles. Hatchery juveniles were also immobile for less time and spent a shorter amount of time than wild juveniles in the fringe of the open‐field arena. Salmo salar size had no effect on the observed behaviour. Overall, this study provides empirical evidence that one generation of hatchery rearing does not change reflex responses associated with threats, whereas antipredator behaviour, typically associated with prior experience, was less developed in hatchery‐reared than in wild individuals.     

A short hatchery history: does it make a difference to aggressiveness in European grayling?

The level of aggressive behaviour in three populations of grayling Thymallus thymallus was lower in the hatchery strains than in the wild strains at the age of 0+ years. Due to similar rearing conditions, genetic divergence of the strains was most likely. As the hatchery fish used were second generation hatchery fish, this suggested that genetic changes in the hatchery can be very rapid. Therefore, it would be beneficial to use the progeny of wild fish for re-introductions. Differences in aggressiveness between the strains still existed at the age of 1+ years, when the strains had been reared under common hatchery conditions for a year. A relatively short period in the hatchery may maintain the original behavioural characteristics of the fish and thus give the best possible basis for survival in the natural environment.     

Differences in the pattern of antipredator behaviour between hatchery‐reared and wild European sea bass juveniles

Shoals of hatchery‐reared and wild sea bass juveniles Dicentrarchus labrax were tested for differences in their antipredator responses towards a potential live predator, the eel Anguilla anguilla . Eight experimental shoals ( i.e . replicates), each composed of 15 individuals from the same stock of juveniles ( i.e . wild or hatchery), were video recorded for 5 min before and after predator exposure. A set of behavioural variables were measured during the pre‐stimulus and stimulus phases of each test and compared between the two groups of replicates. Results showed that in both hatchery‐reared and wild juveniles predator exposure elicited a significant increase in the mean level of shoal cohesiveness and mean shoal distance from the predator, and a significant decrease in the mean shoal distance from the bottom. Shoals of wild juveniles, however, aggregated more quickly and reached higher shoal cohesiveness within the first 20 s of the stimulus period than shoals of hatchery‐reared fish. During this period, the wild fish also reached the highest peak in shoal cohesiveness, which then decreased gradually towards the levels observed before predator exposure. Another component of the antipredator response, the predator inspection behaviour, was fully developed in both wild and hatchery fish. Wild fish, however, tended to inspect the predator at a closer distance than hatchery fish.     

Coordinated evolution of brain size,structure, and eye size in Trinidadian killifish

Brain size, brain architecture, and eye size vary extensively in vertebrates. However, the extent to which the evolution of these components is intricately connected remains unclear. Trinidadian killifish, Anablepsoides hartii, are found in sites that differ in the presence and absence of large predatory fish. Decreased rates of predation are associated with evolutionary shifts in brain size; males from sites without predators have evolved a relatively larger brain and eye size than males from sites with predators. Here, we evaluated the extent to which the evolution of brain size, brain structure, and eye size covary in male killifish. We utilized wild‐caught and common garden‐reared specimens to determine whether specific components of the brain have evolved in response to differences in predation and to determine if there is covariation between the evolution of brain size, brain structure, and eye size. We observed consistent shifts in brain architecture in second generation common garden reared, but not wild caught preserved fish. Male killifish from sites that lack predators exhibited a significantly larger telencephalon, optic tectum, cerebellum, and dorsal medulla when compared with fish from sites with predators. We also found positive connections between the evolution of brain structure and eye size but not between overall brain size and eye size. These results provide evidence for evolutionary covariation between the components of the brain and eye size. Such results suggest that selection, directly or indirectly, acts upon specific regions of the brain, rather than overall brain size, to enhance visual capabilities.     

Effect of hatchery environment on cranial morphology and developmental stability of Atlantic salmon (Salmo salar L.) from North‐West Russia

The study addresses the effect of hatchery rearing on morphological variation and developmental stability of Atlantic salmon parr from North‐West Russia. Totally, we collected nine samples. Four wild samples were collected from each of the rivers Kola, Umba, Keret’ and Shuia. Five samples of hatchery‐reared parr were the first‐generation progeny of wild adults from these rivers reared separately at the four hatcheries (one hatchery was represented by two samples). Ten meristic and 48 morphometric cranial characters were analysed. We studied the morphological divergence between wild and hatchery fishes of the same river of origin. To analyze developmental stability we used fluctuating asymmetry (random deviations from perfect bilateral symmetry). It was found that hatchery‐reared parr significantly differ from wild parr in both meristic characters and the shape of cranial bones. Different hatcheries caused similar effect on morphological variation in all populations. Fluctuating asymmetry in morphometric characters was significantly higher in hatchery fish than in wild from the Shuia River, indicating a higher level of developmental instability. However, wild parr from the Keret’ River had significantly higher fluctuating asymmetry than cultivated parr of the same origin, possible due to a high infection pressure of the parasite Gyrodactylus salaris Malmberg which has led to significant decline of the wild salmon population in this river, or from genetic changes caused by cultivation. The obtained results indicate a notable effect of hatchery environment on Atlantic salmon’s phenotype.     

Effects of the rearing environment on average behaviour and behavioural variation in steelhead

In the context of conservation hatcheries that seek to bolster wild populations by releasing captively-reared fishes into the wild, steelhead Oncorhynchus mykiss were used to test the hypothesis that naturalistic rearing environments promote adaptive behaviour that might otherwise not develop in typical hatchery environments. When comparisons were made among fish reared in barren, structured or structurally variable environments ( i.e. the location of the structure was repositioned every 2–3 days), structure in the rearing environment increased future exploratory behaviour, but only if the structure was stable. Under conditions of high perceived predation risk, the fish no longer exhibited increased exploratory behaviour, suggesting that it is expressed in an adaptive, context-dependant manner. Another concern with hatcheries is that relaxed selection over multiple generations in captivity can increase maladaptive behavioural variation. Compared to rearing in hatchery-typical barren environments, rearing in structured-stable environments decreased behavioural variation. This effect, which occurred during development and did not involve selection, demonstrates a different mechanism for change in behavioural variation in captivity. These experiments show that effects of structure and structural stability occur at the level of both average behaviour and behavioural variation, and suggest that these effects should be considered when fishes are reared in hatcheries for later release into the wild.     

Determination of body shape variation in Irish hatchery‐reared and wild Atlantic salmon

Discriminant function analysis was used to distinguish morphologically between samples of parr, smolts and adult Atlantic salmon Salmo salar from several hatchery and river systems in Ireland. The effect of habitat shift was investigated in Atlantic salmon parr. Parr grown from the eyed‐egg stage with a non‐sibling group in a hatchery environment, came to resemble the mean body shape of their host hatchery Atlantic salmon stock more closely than that of a full sibling group grown at their natal hatchery. Wild Atlantic salmon smolts differed in shape from hatchery‐reared smolts. This difference was less pronounced, but still statistically significant when wild adults were compared with hatchery‐reared adults captured in the coastal drift‐net fishery after a year spent at sea. Rearing conditions had a significant impact on the production and growth of fish body shape. This in turn may have affected adaptability and survivorship of ranched Atlantic salmon in the marine environment.     

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.