Early Marine Migration Patterns of Wild Coastal Cutthroat Trout (Oncorhynchus clarki clarki), Steelhead Trout (Oncorhynchus mykiss), and Their Hybrids

Background

Hybridization between coastal cutthroat trout (Oncorhynchus clarki clarki) and steelhead or rainbow trout (Oncorhynchus mykiss) has been documented in several streams along the North American west coast. The two species occupy similar freshwater habitats but the anadromous forms differ greatly in the duration of marine residence and migration patterns at sea. Intermediate morphological, physiological, and performance traits have been reported for hybrids but little information has been published comparing the behavior of hybrids to the pure species.

Methodology/Principal Findings

This study used acoustic telemetry to record the movements of 52 cutthroat, 42 steelhead x cutthroat hybrids, and 89 steelhead smolts, all wild, that migrated from Big Beef Creek into Hood Canal (Puget Sound, Washington). Various spatial and temporal metrics were used to compare the behavior of the pure species to their hybrids. Median hybrid residence time, estuary time, and tortuosity values were intermediate compared to the pure species. The median total track distance was greater for hybrids than for either cutthroat or steelhead. At the end of each track, most steelhead (80%) were located near or north of the Hood Canal, as expected for this seaward migrating species, whereas most cutthroat (89%) were within 8 kilometers of the estuary. Most hybrids (70%) were detected leaving Hood Canal, though a substantial percentage (20%) remained near the Big Beef Creek estuary. More hybrids (7.5%) than pure cutthroat (4.5%) or steelhead (0.0%) were last detected in the southern reaches of Hood Canal.

Conclusions/Significance

Given the similarity in freshwater ecology between the species, differences in marine ecology may play an important role in maintaining species integrity in areas of sympatry.     

Who are the missing parents? Grandparentage analysis identifies multiple sources of gene flow into a wild population

In order to increase the size of declining salmonid populations, supplementation programmes intentionally release fish raised in hatcheries into the wild. Because hatchery-born fish often have lower fitness than wild-born fish, estimating rates of gene flow from hatcheries into wild populations is essential for predicting the fitness cost to wild populations. Steelhead trout (Oncorhynchus mykiss) have both freshwater resident and anadromous (ocean-going) life history forms, known as rainbow trout and steelhead, respectively. Juvenile hatchery steelhead that 'residualize' (become residents rather than go to sea as intended) provide a previously unmeasured route for gene flow from hatchery into wild populations. We apply a combination of parentage and grandparentage methods to a three-generation pedigree of steelhead from the Hood River, Oregon, to identify the missing parents of anadromous fish. For fish with only one anadromous parent, 83% were identified as having a resident father while 17% were identified as having a resident mother. Additionally, we documented that resident hatchery males produced more offspring with wild anadromous females than with hatchery anadromous females. One explanation is the high fitness cost associated with matings between two hatchery fish. After accounting for all of the possible matings involving steelhead, we find that only 1% of steelhead genes come from residualized hatchery fish, while 20% of steelhead genes come from wild residents. A further 23% of anadromous steelhead genes come from matings between two resident parents. If these matings mirror the proportion of matings between residualized hatchery fish and anadromous partners, then closer to 40% of all steelhead genes come from wild trout each generation. These results suggest that wild resident fish contribute substantially to endangered steelhead 'populations' and highlight the need for conservation and management efforts to fully account for interconnected Oncorhynchus mykiss life histories.     

Seasonal foraging and piscivory by sympatric wild and hatchery-reared steelhead from an integrated hatchery program

We compared the diet of hatchery-reared steelhead produced from an integrated hatchery program as emigrating spring smolts and non-migrating hatchery residuals to their sympatric wild counterparts. Our results suggest that there is a potential for hatchery fish to affect wild steelhead populations due to dietary overlap and subyearling salmonid predation; however, relative ecological risk did not increase as steelhead delayed or forwent emigration. Predation by hatchery smolts was related to release timing, but not experience with native fish. Diet composition appears to be more strongly affected by seasonal and yearly differences in prey abundance and presence rather than differences in rearing environments. Hatchery and wild steelhead showed small but important foraging differences. Hatchery smolts did not consume as many salmonids as wild fish and hatchery residuals showed relatively stronger surface oriented feeding behavior than wild parr. Because most hatchery smolts emigrated shortly after release and the overall number of residuals in the study creek was low, we speculate that in this case there is low dietary and predatory-based risk of hatchery steelhead in Abernathy Creek negatively impacting wild salmonids.     

Development of natural growth regimes for hatchery-reared steelhead to reduce residualism, fitness loss, and negative ecological interactions

Wild steelhead (Oncorhynchus mykiss) typically spend two or more years in freshwater before migrating to sea, but hatchery steelhead are almost ubiquitously released as yearlings. Their large size at release coupled with life history pathways that include both male and female maturation in freshwater present ecological risks different from those posed by hatchery populations of Pacific salmon. Yearling hatchery reared steelhead that fail to attain minimum thresholds for smoltification or exceed thresholds for male maturation tend to ‘residualize’ (i.e., remain in freshwater). Residuals pose ecological risks including size-biased interference competition and predation on juvenile salmon and trout. Three hatchery populations of steelhead in Hood Canal, WA were reared under growth regimes designed to produce a more natural age at smoltification (age-2) to aid in rebuilding their respective natural populations. Mean smolt sizes and size variability at age-2 were within the range of wild smolts for two of the three populations. The third population reared at a different facility under similar temperatures exhibited high growth rate variability and high male maturation rates (20% of all released fish). Experimentally comparing age-1 and age-2 smolt programs will help identify optimal rearing strategies to reduce the genetic risk of domestication selection and reduce residualism rates and associated negative ecological effects on natural populations. Investigations of Winthrop National Fish Hatchery summer-run steelhead will measure a) selection on correlated behavioral traits (‘behavioral syndromes’), b) degree of smoltification, c) changes in hormones that regulate gonad growth at key developmental stages, and d) conduct extensive post-release monitoring of fish reared under each growth regime.     

Riverine, estuarine and marine migratory behaviour and physiology of wild and hatchery-reared coho salmon Oncorhynchus kisutch (Walbaum) smolts descending the Campbell River, BC, Canada

Eighty coho salmon Oncorhynchus kisutch smolts (40 wild and 40 hatchery-reared) were surgically implanted with acoustic transmitters and released into the Quinsam River over 2 days. Differences in physiology, travel time and migratory behaviour were examined between wild and hatchery-reared fish. In addition, tagged and control fish of both wild and hatchery-reared stock were raised for 3 months following surgery to compare survival and tag retention. Detection ranges of the acoustic receivers were tested in the river, estuary and ocean in a variety of flow conditions and tide levels. Receivers were placed in the river, estuary and up to 50 km north and south from the river mouth in the marine environment. Wild smolts were significantly smaller by mass, fork length and condition factor than hatchery-reared smolts and exhibited significantly higher levels of sodium, potassium and chloride in their blood plasma than hatchery-reared smolts. The gill Na⁺K⁺-ATPase activity was also significantly higher in the wild coho smolts at the time of release. Ninety-eight per cent of wild and 80% of hatchery-reared fish survived to the estuary, 8 km downstream of the release site. No difference was found in migration speed, timing or survival between smolts released during daylight and those released after dark. Wild smolts, however, spent less time in the river and estuary, and as a result entered the ocean earlier than hatchery-reared smolts. Average marine swimming speeds for wild smolts were double those of their hatchery-reared counterparts. While hatchery smolts dispersed in both a northward and southward direction upon entering the marine environment, the majority of wild smolts travelled north from the Campbell River estuary. The wild coho salmon smolts were more physiologically fit and ready to enter sea water than the hatchery-reared smolts, and as a result had higher early survival rates and swimming speeds.     

Recent Salmon Declines: A Result of Lost Feeding Opportunities Due to Bad Timing?

As the timing of spring productivity blooms in near-shore areas advances due to warming trends in global climate, the selection pressures on out-migrating salmon smolts are shifting. Species and stocks that leave natal streams earlier may be favoured over later-migrating fish. The low post-release survival of hatchery fish during recent years may be in part due to static release times that do not take the timing of plankton blooms into account. This study examined the effects of release time on the migratory behaviour and survival of wild and hatchery-reared coho salmon (Oncorhynchus kisutch) using acoustic and coded-wire telemetry. Plankton monitoring and near-shore seining were also conducted to determine which habitat and food sources were favoured. Acoustic tags (n = 140) and coded-wire tags (n = 266,692) were implanted into coho salmon smolts at the Seymour and Quinsam Rivers, in British Columbia, Canada. Differences between wild and hatchery fish, and early and late releases were examined during the entire lifecycle. Physiological sampling was also carried out on 30 fish from each release group. The smolt-to-adult survival of coho salmon released during periods of high marine productivity was 1.5- to 3-fold greater than those released both before and after, and the fish''s degree of smoltification affected their downstream migration time and duration of stay in the estuary. Therefore, hatchery managers should consider having smolts fully developed and ready for release during the peak of the near-shore plankton blooms. Monitoring chlorophyll a levels and water temperature early in the spring could provide a forecast of the timing of these blooms, giving hatcheries time to adjust their release schedule.     

Differential Reproductive Success of Sympatric, Naturally Spawning Hatchery and Wild Steelhead, Oncorhynchus mykiss

Hatchery propagation of salmonids has been practiced in western North America for over a century. However, recent declines in wild salmon abundance and efforts to mitigate these declines through hatcheries have greatly increased the relative abundance of fish produced in hatcheries. The over-harvest of wild salmon by fishing mixed hatchery and wild stocks has been of concern for many years but genetic interactions between populations, such as hybridization, introgression and outbreeding depression, may also compromise the sustainability of wild populations. Our goal was to examine whether a newly established hatchery population of steelhead trout successfully reproduced in the wild and to compare their rate of reproductive success to that of sympatrically spawning native steelhead. We used eight microsatellite loci to create allele frequency profiles for baseline hatchery and wild populations and assigned the smolt (age 2) offspring of this parental generation to a population of origin. Adults originating from a generalized hatchery stock artificially selected for early return and spawning date were successful at reproducing in Forks Creek, Washington. Although hatchery females (N = 90 and 73 in the two consecutive years of the study) produced offspring that survived to emigrate as smolts, they produced only 4.4–7.0% the number produced per wild female (N = 11 and 10). This deficit in reproductive success implies that the proportion of hatchery genes in the mixed population may diminish since deliberate releases into the river have ceased. This hypothesis is being tested in a long-term study at Forks Creek.     

Interactions between endangered wild and hatchery salmonids: can the pitfalls of artificial propagation be avoided in small coastal streams?

Hatchery and wild juvenile populations of steelhead Oncorhynchus mykiss and coho salmon Oncorhynchus kisutch , in a small coastal watershed in central California, were sampled throughout the year in a stream and at a hatchery. Both species grew faster in captivity than in the wild. Hatchery fish of both species had elevated gill Na⁺, K⁺‐ATPase activity, and thus were ready to enter sea water when planted during the wild fish migration. Downstream migrant trapping and stream surveys indicated that hatchery smolts went to sea soon after planting, consequently avoiding the effects of competition and predation that commonly occur when hatchery‐bred juveniles are released. Adult steelhead were also sampled throughout the watershed. The return of hatchery steelhead was highly synchronized with that of wild steelhead, indicating that hatchery propagation had no adverse effects on the timing of the run. A disproportionate number of hatchery steelhead returned to the tributary where the hatchery was located, despite being planted throughout the watershed. Hatchery steelhead did not differ in mean age or size from wild steelhead. Observations of spawning indicated that hatchery and wild steelhead interbreed. Competition for mates or spawning substratum was rarely observed between hatchery and wild steelhead. Many of the problems commonly associated with artificial propagation can be avoided in small coastal watersheds when wild broodstock are used and fish are released as smolts.     

Effective size of a wild salmonid population is greatly reduced by hatchery supplementation

Many declining and commercially important populations are supplemented with captive-born individuals that are intentionally released into the wild. These supplementation programs often create large numbers of offspring from relatively few breeding adults, which can have substantial population-level effects. We examined the genetic effects of supplementation on a wild population of steelhead (Oncorhynchus mykiss) from the Hood River, Oregon, by matching 12 run-years of hatchery steelhead back to their broodstock parents. We show that the effective number of breeders producing the hatchery fish (broodstock parents; N(b)) was quite small (harmonic mean N(b)=25 fish per brood-year vs 373 for wild fish), and was exacerbated by a high variance in broodstock reproductive success among individuals within years. The low N(b) caused hatchery fish to have decreased allelic richness, increased average relatedness, more loci in linkage disequilibrium and substantial levels of genetic drift in comparison with their wild-born counterparts. We also documented a substantial Ryman-Laikre effect whereby the additional hatchery fish doubled the total number of adult fish on the spawning grounds each year, but cut the effective population size of the total population (wild and hatchery fish combined) by nearly two-thirds. We further demonstrate that the Ryman-Laikre effect is most severe in this population when (1) >10% of fish allowed onto spawning grounds are from hatcheries and (2) the hatchery fish have high reproductive success in the wild. These results emphasize the trade-offs that arise when supplementation programs attempt to balance disparate goals (increasing production while maintaining genetic diversity and fitness).     

Strategies for reducing the ecological risks of hatchery programs: Case studies from the Pacific Northwest

The Pacific Northwest state and federal agencies and tribes that operate salmon and steelhead (Oncorhynchus sp.) hatcheries are authorized to develop and implement strategies to reduce the risks the programs pose to wild fish populations. This paper reviews five case studies from the states of Oregon and Washington, USA, where agencies and tribes have implemented or proposed programs that were intended to reduce ecological risks due to hatchery programs. The case studies are for Oregon coho salmon, Select Area terminal fisheries programs for Chinook and coho salmon in the lower Columbia River, Hood Canal chum salmon in Puget Sound Washington, Siletz River steelhead on the Oregon coast, and Okanogan River Chinook salmon in eastern Washington. The five case studies address a diversity of management objectives and species. They demonstrate some of the science and risk reduction strategies used to alleviate the ecological effects of hatcheries, and they document some of the results and outcomes of taking action. Elements of four of the case studies have been in place for nearly 20 years. The available science and the conservation ethic toward hatchery programs evolved significantly over this period, and management decisions and strategies have been influenced by public policy as well as by scientific information. Therefore the case studies also document some of the history, the evolution of ideas, the uncertainty, and the political controversy associated with the management of this risk factor. The paper concludes with six principles to help guide the development of future risk reduction programs.     

Do hatchery-reared sea urchins pose a threat to genetic diversity in wild populations?

In salmonids, the release of hatchery-reared fish has been shown to cause irreversible genetic impacts on wild populations. However, although responsible practices for producing and releasing genetically diverse, hatchery-reared juveniles have been published widely, they are rarely implemented. Here, we investigated genetic differences between wild and early-generation hatchery-reared populations of the purple sea urchin Paracentrotus lividus (a commercially important species in Europe) to assess whether hatcheries were able to maintain natural levels of genetic diversity. To test the hypothesis that hatchery rearing would cause bottleneck effects (that is, a substantial reduction in genetic diversity and differentiation from wild populations), we compared the levels and patterns of genetic variation between two hatcheries and four nearby wild populations, using samples from both Spain and Ireland. We found that hatchery-reared populations were less diverse and had diverged significantly from the wild populations, with a very small effective population size and a high degree of relatedness between individuals. These results raise a number of concerns about the genetic impacts of their release into wild populations, particularly when such a degree of differentiation can occur in a single generation of hatchery rearing. Consequently, we suggest that caution should be taken when using hatchery-reared individuals to augment fisheries, even for marine species with high dispersal capacity, and we provide some recommendations to improve hatchery rearing and release practices. Our results further highlight the need to consider the genetic risks of releasing hatchery-reared juveniles into the wild during the establishment of restocking, stock enhancement and sea ranching programs.     

The effect of hatchery release strategy on marine migratory behaviour and apparent survival of Seymour River steelhead smolts (Oncorhynchus mykiss)

Early marine migratory behaviour and apparent survival of hatchery-reared Seymour River steelhead (Oncorhynchus mykiss) smolts was examined over a four year period (2006-2009) to assess the impact of various management strategies on improving early marine survival. Acoustically tagged smolts were released to measure their survival using estuary and coastal marine receivers forming components of the Pacific Ocean Shelf Tracking (POST) array. Early marine survival was statistically indistinguishable between releases of summer run and winter run steelhead races, night and day releases, and groups released 10 days apart. In 2009, the survival of summer run steelhead released into the river was again trialed against groups released directly into the ocean at a distance from the river mouth. Apparent survival was improved significantly for the ocean released groups. The health and physiological status of the various release groups were monitored in years 2007-2009, and results indicate that the fish were in good health, with no clinical signs of disease at the time of release. The possibility of a disease event contributing to early marine mortality was further examined in 2009 by vaccinating half of the released fish against common fish diseases (vibriosis, furunculosis). The results suggest that marine survival may be enhanced using this approach, although not to the extent observed when the smolts were transported away from the river mouth before release. In summary, direct experimental testing of different release strategies using the POST array to measure ocean survival accelerated the scientific process by allowing rapid collection of data which enabled the rejection of several existing theories and allowed tentative identification of several new alternative approaches that might improve early marine survival of Seymour River steelhead.     

Restitution and genetic differentiation of salmon populations in the southern Baltic genotyped with the Atlantic salmon 7K SNP array

Background

Native populations of Atlantic salmon in Poland, from the southern Baltic region, became extinct in the 1980s. Attempts to restitute salmon populations in Poland have been based on a Latvian salmon population from the Daugava river. Releases of hatchery reared smolts started in 1986, but to date, only one population with confirmed natural reproduction has been observed in the Slupia river. Our aim was to investigate the genetic differentiation of salmon populations in the southern Baltic using a 7K SNP (single nucleotide polymorphism) array in order to assess the impact of salmon restitution in Poland.

Methods

One hundred and forty salmon samples were collected from: the Polish Slupia river including wild salmon and individuals from two hatcheries, the Swedish Morrum river and the Lithuanian Neman river. All samples were genotyped using an Atlantic salmon 7K SNP array. A set of 3218 diagnostic SNPs was used for genetic analyses.

Results

Genetic structure analyses indicated that the individuals from the investigated populations were clustered into three groups i.e. one clade that included individuals from both hatcheries and the wild population from the Polish Slupia river, which was clearly separated from the other clades. An assignment test showed that there were no stray fish from the Morrum or Neman rivers in the sample analyzed from the Slupia river. Global F_ST over polymorphic loci was high (0.177). A strong genetic differentiation was observed between the Lithuanian and Swedish populations (F_ST = 0.28).

Conclusions

Wild juvenile salmon specimens that were sampled from the Slupia river were the progeny of fish released from hatcheries and, most likely, were not progeny of stray fish from Sweden or Lithuania. Strong genetic differences were observed between the salmon populations from the three studied locations. Our recommendation is that future stocking activities that aim at restituting salmon populations in Poland include stocking material from the Lithuanian Neman river because of its closer geographic proximity.

Electronic supplementary material

The online version of this article (doi:10.1186/s12711-015-0121-9) contains supplementary material, which is available to authorized users.     

Feeding of wild and hatchery reared Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis> L.) smolts during downstream migration

In general, hatchery salmonid smolts experience higher mortality during migration than wild smolts, which is suggested to be due to domestication effects and that hatchery fish lack experience of the natural environment. However, possible differences in feeding during smolt migration between hatchery and wild smolts have rarely been addressed. We compared the number of feeding smolts and stomach fullness among wild Atlantic salmon smolts, hatchery-reared smolts released as 1-year-old parr, and hatchery-reared smolts released as 2-year-old smolts during their descent to sea in River Tornionjoki. In addition, estimations of prey selection among the smolt groups were conducted. A high proportion of wild smolts and smolts stocked as parr actively fed during the smolt migration. A lower proportion of smolts stocked as smolts was feeding and their stomach fullness were much reduced in comparison with the two other groups. The study also indicated that the feeding of migrating smolts is selective rather than opportunistic. In conclusion, this study suggests that stocked 2-year-old smolts may enter sea with an inferior foraging behaviour and it is a possibility that this may contribute to the observed low post-smolt survival in the Baltic Sea.     

Survival and migration patterns of naturally and hatchery-reared Atlantic salmon (Salmo salar) smolts in a Lake Ontario tributary using acoustic telemetry

1. Atlantic salmon (Salmo salar) smolts are often stocked into rivers to supplement natural reproduction, yet hatchery-reared fish have lower survival compared to wild conspecifics. However, few studies have assessed riverine migratory performance and survival differences in hatchery and wild smolts, or more specifically naturally reared smolts (hatchery fish released earlier as parr), particularly in rivers with weirs which may further reduce survival.
2. Using acoustic telemetry, including a subset of fish with novel transmitters that identify predation events, we assessed survival and migration patterns of hatchery- (2017: n = 32; 2018: n = 30) and naturally reared Atlantic salmon smolts (2017: n = 8; 2018: n = 30) in a Lake Ontario tributary with two weirs to better understand their ecology and assess the influence of environmental parameters on migration.
3. Naturally reared smolts were 13.9 times more likely to survive than hatchery-reared smolts and mark–recapture models indicated that weirs did not reduce survival for either group. Survival per km was lowest at the release site, indicating pre-migration mortality, and specifically high stocking-related mortality of hatchery-reared smolts. Speed and times of day fish migrated (i.e. migratory performance) did not vary by rearing group, suggesting that the high mortality of hatchery-reared smolts may be due to other factors related to hatchery and stocking operations. Overall mean (± SD) migration speed for smolts was 0.70 ± 0.39 km/hr and movements occurred significantly more frequently at night (18:00–06:00).
4. Smolts were detected in Lake Ontario after they left the river; however, the array in Lake Ontario was not conducive to providing much detail regarding movement patterns. There was no predation of the two predation tags detected in Lake Ontario, indicating that movements were made by smolts and not predators.
5. With ongoing restoration efforts of Atlantic salmon in Lake Ontario, it was important to understand the smolt migration patterns and success of the stocked fish. Our findings of similar migratory performance yet different relative survival of hatchery- and naturally reared smolts help inform management with regards to stocking strategies that could improve Atlantic salmon reintroduction success.

     

Differences in Lateral Line Morphology between Hatchery- and Wild-Origin Steelhead

Despite identification of multiple factors mediating salmon survival, significant disparities in survival-to-adulthood among hatchery- versus wild-origin juveniles persist. In the present report, we explore the hypothesis that hatchery-reared juveniles might exhibit morphological defects in vulnerable mechanosensory systems prior to release from the hatchery, potentiating reduced survival after release. Juvenile steelhead (Oncorhynchus mykiss) from two different hatcheries were compared to wild-origin juveniles on several morphological traits including lateral line structure, otolith composition (a proxy for auditory function), and brain weight. Wild juveniles were found to possess significantly more superficial lateral line neuromasts than hatchery-reared juveniles, although the number of hair cells within individual neuromasts was not significantly different across groups. Wild juveniles were also found to possess primarily normal, aragonite-containing otoliths, while hatchery-reared juveniles possessed a high proportion of crystallized (vaterite) otoliths. Finally, wild juveniles were found to have significantly larger brains than hatchery-reared juveniles. These differences together predict reduced sensitivity to biologically important hydrodynamic and acoustic signals from natural biotic (predator, prey, conspecific) and abiotic (turbulent flow, current) sources among hatchery-reared steelhead, in turn predicting reduced survival fitness after release. Physiological and behavioral studies are required to establish the functional significance of these morphological differences.     

Impact of parasites on salmon recruitment in the Northeast Atlantic Ocean

Parasites may have large effects on host population dynamics, marine fisheries and conservation, but a clear elucidation of their impact is limited by a lack of ecosystem-scale experimental data. We conducted a meta-analysis of replicated manipulative field experiments concerning the influence of parasitism by crustaceans on the marine survival of Atlantic salmon (Salmo salar L.). The data include 24 trials in which tagged smolts (totalling 283 347 fish; 1996–2008) were released as paired control and parasiticide-treated groups into 10 areas of Ireland and Norway. All experimental fish were infection-free when released into freshwater, and a proportion of each group was recovered as adult recruits returning to coastal waters 1 or more years later. Treatment had a significant positive effect on survival to recruitment, with an overall effect size (odds ratio) of 1.29 that corresponds to an estimated loss of 39 per cent (95% CI: 18–55%) of adult salmon recruitment. The parasitic crustaceans were probably acquired during early marine migration in areas that host large aquaculture populations of domesticated salmon, which elevate local abundances of ectoparasitic copepods—particularly Lepeophtheirus salmonis. These results provide experimental evidence from a large marine ecosystem that parasites can have large impacts on fish recruitment, fisheries and conservation.     

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.     

Effects of a single event of close inbreeding on growth and survival in steelhead

For two populations of Alaskan steelhead (Oncorhynchus mykiss) of common ancestry we evaluated effects of inbreeding in second-generation descendants of wild fish by comparing progeny of full-sibling matings to those of non-inbred controls to determine if a single event of close inbreeding has significant effects on survival and growth in captivity or the wild. In captivity, both survival and size were highly variable between inbred and control types within each line and among the five broods during five periods of freshwater culture. However, no consistent patterns of inbreeding enhancement or depression between types within lines across years were evident. In contrast, in the wild marine environment, 34 of 34 pairwise comparisons between inbred and control types in body size of returning adults after 2 or 3 years at liberty in the ocean were consistent with inbreeding depression with significant inbreeding depression varying from 2.9% for female length to 20.0% for female weight. Survival of marked juveniles (smolts) to adults in the wild marine environment was consistently and significantly lower in inbred types for both lines, for an average inbreeding depression of 78.8%. The results underscore the potential problems that can arise from using protective culture technologies, including captive broodstocks, to supplement endangered populations, and they highlight the genetic hazards that can be faced by small wild populations. This study demonstrates that high natural mortality or selection increases the amount of inbreeding depression detected in survival. Inbreeding effects on survival and growth in captivity can be poor indicators of survival and growth in a wild marine environment.