Social status and growth rates as determinants of site attachment in juvenile Atlantic salmon

This paper describes an experimental study of the effects of food supply, growth rates and social interactions on homing by juvenile Atlantic salmon Salmo salar in response to displacement. Groups of five fish were housed in a section of an artificial stream and given either rations allowing maximum growth (the rich condition) or 0·1 of this amount (the poor condition); daily specific growth rates were significantly higher in the rich condition. After a 6-day settlement period, the fish were captured, displaced downstream and their movements recorded over the next 3 h. Prior to displacement, the fish showed a high degree of site fidelity and high levels of aggression. Dominant fish and those with stronger site attachment grew faster prior to displacement, these effects being independent. Following displacement, 24% of all fish returned to their previously favoured site and stayed there, 23% returned home initially, but subsequently moved on, 5% settled in a new site and 49% failed to move. The distribution of responses was identical for the rich and poor conditions, but fish that homed were dominant and had grown faster during the pre-displacement period.     

Changes in serum thyroxine levels during smolting in captive and wild Atlantic salmon, Salmo salar L.

On a commercial fish farm in Western Scotland, smolting Atlantic salmon, Salmo salar L., showed no major changes in serum thyroxine (T4) levels in the spring. In contrast, serum T4 levels in smolting, wild salmon, trapped as they left the Girnock Burn, Aberdeenshire, varied markedly over this period. Serum T4 levels were correlated with the stream discharge rate pertaining as the fish left the stream. Levels in migrating, smolting fish were higher than those in fish captured above the fish-trap by electro-fishing. In a stream channel, serum T4 levels in smolting fish caught previously in the fish-trap were rapidly elevated by exposure to increased water-flow, whereas levels in fish previously captured above the trap remained substantially unchanged. Serum T4 levels in salmon parr leaving the stream in autumn were generally lower than those in smolting fish leaving in spring and were not demonstrably affected by stream discharge rate.     

Effects of Emergence Time and Early Social Rearing Environment on Behaviour of Atlantic Salmon: Consequences for Juvenile Fitness and Smolt Migration

Consistent individual differences in behaviour have been well documented in a variety of animal taxa, but surprisingly little is known about the fitness and life-history consequences of such individual variation. In wild salmonids, the timing of fry emergence from gravel spawning nests has been suggested to be coupled with individual behavioural traits. Here, we further investigate the link between timing of spawning nest emergence and behaviour of Atlantic salmon (Salmo salar), test effects of social rearing environment on behavioural traits in fish with different emergence times, and assess whether behavioural traits measured in the laboratory predict growth, survival, and migration status in the wild. Atlantic salmon fry were sorted with respect to emergence time from artificial spawning nest into three groups: early, intermediate, and late. These emergence groups were hatchery-reared separately or in co-culture for four months to test effects of social rearing environment on behavioural traits. Twenty fish from each of the six treatment groups were then subjected to three individual-based behavioural tests: basal locomotor activity, boldness, and escape response. Following behavioural characterization, the fish were released into a near-natural experimental stream. Results showed differences in escape behaviour between emergence groups in a net restraining test, but the social rearing environment did not affect individual behavioural expression. Emergence time and social environment had no significant effects on survival, growth, and migration status in the stream, although migration propensity was 1.4 to 1.9 times higher for early emerging individuals that were reared separately. In addition, despite individuals showing considerable variation in behaviour across treatment groups, this was not translated into differences in growth, survival, and migration status. Hence, our study adds to the view that fitness (i.e., growth and survival) and life-history predictions from laboratory measures of behaviour should be made with caution and ideally tested in nature.     

Movements of brown trout, Salmo trutta, and juvenile Atlantic salmon, Salmo salar, in a coastal stream in northern Norway

Movements of resident brown trout (age 2+ to 9+ years) and young Atlantic salmon (age 1+), stocked as fry, were studied in July, August and September in a coastal stream in northern Norway. Between 85 and 89% of the brown trout were recaptured in the study area (346m, 1326m²) within 45m of their release point throughout the investigation period. Most specimens had moved less than 150m. Trout movements were related to local variation in density. Trout occupying those sections of stream with the lowest fish densities (5.3–10.9 fish 100m^?2) had a significantly lower movement rate than fish from sections with densities between 13.7 and 31.5 fish 100m^?2. Trout that moved from their marking section were significantly larger than specimens that did not leave their original site. There was a significant correlation between permanence of station each month and the mean water level that month. The majority of the trout (47%) were caught at undercut stream banks or at sites in the proximity of these. A decrease in water level during the study period resulted in a high loss (36%) of such habitat, probably forcing some individuals to move. The recapture data indicate that the trout population consists of one small (c. 15–20%) mobile, and one large sedentary component. Young salmon displayed high station permanence in July and August (93% and 96%). However, in the autumn they exhibited a significant downstream movement, and only 73% were recaptured within their release section. This movement was significantly higher for larger specimens, and is thought to occur because of a pre-winter change in habitat, initiated by a decline in stream temperature. In contrast to trout, salmon in sections containing the lowest densities (22.0–25.0 fish 100m^?2) did not show significantly lower movement rates when compared with salmon at higher densities (32.2–46.3 and 51.8–60.6 fish 100m^?2). The spatial distribution of young salmon indicated the formation of territorial mosaics over the stream bed, which are thought to reduce intraspecific competition.     

Effects of temperature and growth hormone on individual growth trajectories of wild‐type and transgenic coho salmon Oncorhynchus kisutch

In this study, individual growth patterns of wild‐type and growth‐enhanced coho salmon Oncorhynchus kisutch at 8, 12 and 16° C water temperature were followed. Despite large differences among individuals in growth rates, there was generally little variation in the shape of the growth curves among O. kisutch individuals of both genotypes and at all temperatures. Typically, individuals that were relatively large initially were also relatively large at the end of the growth period. The limitation in variation was more pronounced in the growth‐enhanced O. kisutch than in the wild type, where the relative size of some individuals reared at 12 and 8° C changed by the end of the trial. As a warmer temperature seems to decrease the plasticity of growth trajectories in wild‐type fish, it is possible that global warming will influence the ability of wild fish to adapt their growth to changing conditions.     

Effects of long-term cortisol treatments on gonadal development, sex steroids levels and ovarian cortisol content in cultured great sturgeon Huso huso

The objective of this study was to examine the effect of cortisol implantations on gonadal development, sex steroid levels, and ovarian cortisol content in cultured great sturgeon Huso huso. Three groups of 5 fish for each treatment were considered. The experimental groups included: control (capsules containing cocoa butter alone), low cortisol (C(5); 5mg cortisol/kg body mass+cocoa butter) and, high cortisol (C(50); 50mg cortisol/kg body mass+cocoa butter). The capsules containing hormones and cocoa butter were intraperitoneally implanted into 3-year-old female fish at pre-vitellogenic stage (mean initial body mass 6809.7 ± 73 g) every 6 weeks over a 6-month period from January to June. The serum levels of cortisol, glucose, cholesterol and sex steroids (testosterone and 17β-estradiol) were determined at the initial time and three weeks after each implantation. Oocyte histological characteristics (the diameter and area of the oocyte, the diameter and area of the nucleus and the ratio of the nucleus area to the oocyte area) were measured at the end of the experiment and compared to those at the initial time. Ovarian cortisol content was measured at the end of the experiment. The results showed that serum cortisol levels varied in a dose-independent manner, so that the highest cortisol concentrations were observed in C(5)-treated fish throughout the experiment. Serum glucose levels were significantly higher in cortisol-treated groups than those in the control group. The high dose of cortisol elicited a significant constant increase in serum cholesterol concentrations. Fish implanted with the high cortisol dose showed significant declines in serum testosterone and 17β-estradiol concentrations throughout the experiment. No significant differences were found in oocyte histological characteristics among experimental groups. The cortisol implants elicited a dose-dependent increase in ovarian cortisol content. At the end of trial, body-growth indices were the lowest in C(50)-implanted fish, while the low cortisol dose had no effect on growth relative to the controls. These results indicated that chronic stress induced by cortisol implantation in great sturgeon suppressed gonadal steroidogenesis and somatic growth but had no effect on ovarian growth and development.     

Growth and survival of Atlantic salmon (Salmo salar) in Llyn Dwythwch, North Wales

SUMMARY. Atlantic salmon fry have been annually stocked into Llyn Dwythwch, North Wales, since 1969, in an attempt to increase the natural stocks of the area. The growth and survival of 1- and 2-year-old salmon were investigated, and compared with that of other lake-reared populations and also with salmon in the natural stream environment. Lake-reared salmon follow the same patterns of slow and rapid growth as found for river fish, but the growth rate was superior in the former. The variation in length – weight relationship with age and sex was investigated. Survival rates in general compared favourably with the survival in rivers, with high mortality rates of salmon in Llyn Dwythwch resulting from predation at spring stocking by the resident brown trout. This was later avoided by stocking larger fish in the autumn.     

Testing predictions of the critical period for survival concept using experiments with stocked Atlantic salmon

Two separate field experiments were performed in the U.S.A. and Norway with stocked Atlantic salmon Salmo salar . In the Norwegian experiment, the offspring of early‐spawning fish which had larger eggs and emerged a few days before offspring of later spawning fish had consistently higher survival rates. In the U.S.A. experiment, stream sections with higher proportions of favourable foraging locations during the critical period (the transition from dependence on maternally‐derived yolk reserves to independent feeding) had lower loss rates of fish stocked as unfed fry. These results provide support for the critical period concept (CPC) in Atlantic salmon, underscores the utility of a manipulative approach to achieve further advances in knowledge of Atlantic salmon ecology and provide additional guidance to management and restoration. A mechanistic, conceptual model for density dependence is presented to identify important knowledge gaps that remain to further evaluate the importance of the CPC for Atlantic salmon population regulation.     

Stocking efficiency and the effects of diet preconditioning on the post-release adaptation of hatchery-reared juveniles of Atlantic salmon (Salmo salar L.) in an Atlantic temperate stream

This study explores the stocking efficiency of Atlantic salmon juveniles in a 4th order stream located at the southern limit of its Eastern Atlantic distribution. Moreover, with the idea of implementing new protocols and methods to improve ??traditional?? hatchery practices and rear a more ??wild?? like fish, the study examines the response of this species to natural diet preconditioning prior to release. The field study indicated that most juveniles disappeared from the stocked reaches within two months of their release, resulting in a recapture rate after a year of 0.0?C1.1?%. There were no differences in stocked juvenile??s density along the stream during the first months following stocking. The preconditioning experiments showed that juveniles of Atlantic salmon suffered a significant decrease in their condition factor when diet was changed from artificial pellets to live preys, either if the change occurred before (i.e., in a manipulative experiment) or after their release in the stream. Although weight, length, condition factor and specific growth rates reached by juveniles at the end of the rearing period were higher in individuals fed on pellets than in those fed on macroinvertebrates, differences disappeared only a week after their release in the stream. These results may have implications regarding the stocking success when releasing hatchery-reared Atlantic salmons in suboptimal environments. In particular, the study indicates that stocked salmon adaptation in the selected stream may not be improved by short conditioning periods to natural diet prior to their release, while we discuss potential explanations for the observed results.     

Individual space use strategies of wild juvenile Atlantic salmon

Movements of 60 stream-dwelling wild Atlantic salmon Salmo salar (97–118 mm), each tagged with a passive integrated transponder, were monitored during four trials in an enclosed section (24 m long, 45·1 m² total area) of a stream at a range of densities (four, eight, 16 and 32 fish per enclosure). Patterns of space use differed markedly between individuals, with 80% of fish establishing home ranges within 8 days of introduction to the enclosure (settlers) and the remainder continuing to move throughout the length of the enclosure (non-settlers). Although aggressive interactions were quite frequent and dominant fish were observed chasing subordinates, there was considerable overlap of home ranges of settlers at all densities; this was the case even at lower densities at which only a fraction of the enclosure was used by the fish. Thus, rather than adopting fixed territories, the salmon showed a high level of space sharing. Individual fish used the same local area in different ways, ranging from highly localized feeding on drifting food items to a wider-ranging strategy of specialising on benthic food. Among the fish that settled absolute growth rates were inversely related to body size, and at high densities fish lost weight. These results suggest that space use in wild juvenile salmon is more complex than a mosaic of territories, that salmon demonstrate significant variability in individual space use patterns, and that large fish may suffer disproportionately when populations are at the carrying capacity of their environment.     

Alternate Directed Anthropogenic Shifts in Genotype Result in Different Ecological Outcomes in Coho Salmon Oncorhynchus kisutch Fry

Domesticated and growth hormone (GH) transgenic salmon provide an interesting model to compare effects of selected versus engineered phenotypic change on relative fitness in an ecological context. Phenotype in domestication is altered via polygenic selection of traits over multiple generations, whereas in transgenesis is altered by a single locus in one generation. These established and emerging technologies both result in elevated growth rates in culture, and are associated with similar secondary effects such as increased foraging, decreased predator avoidance, and similar endocrine and gene expression profiles. As such, there is concern regarding ecological consequences should fish that have been genetically altered escape to natural ecosystems. To determine if the type of genetic change influences fitness components associated with ecological success outside of the culture environments they were produced for, we examined growth and survival of domesticated, transgenic, and wild-type coho salmon fry under different environmental conditions. In simple conditions (i.e. culture) with unlimited food, transgenic fish had the greatest growth, while in naturalized stream tanks (limited natural food, with or without predators) domesticated fish had greatest growth and survival of the three fish groups. As such, the largest growth in culture conditions may not translate to the greatest ecological effects in natural conditions, and shifts in phenotype over multiple rather than one loci may result in greater success in a wider range of conditions. These differences may arise from very different historical opportunities of transgenic and domesticated strains to select for multiple growth pathways or counter-select against negative secondary changes arising from elevated capacity for growth, with domesticated fish potentially obtaining or retaining adaptive responses to multiple environmental conditions not yet acquired in recently generated transgenic strains.     

Spatial and trophic overlap of marked and unmarked Columbia River Basin spring Chinook salmon during early marine residence with implications for competition between hatchery and naturally produced fish

Ecological interactions between natural and hatchery juvenile salmon during their early marine residence, a time of high mortality, have received little attention. These interactions may negatively influence survival and hamper the ability of natural populations to recover. We examined the spatial distributions and size differences of both marked (hatchery) and unmarked (a high proportion of which are natural) juvenile Chinook salmon in the coastal waters of Oregon and Washington from May to June 1999–2009. We also explored potential trophic interactions and growth differences between unmarked and marked salmon. Overlap in spatial distribution between these groups was high, although catches of unmarked fish were low compared to those of marked hatchery salmon. Peak catches of hatchery fish occurred in May, while a prolonged migration of small unmarked salmon entered our study area toward the end of June. Hatchery salmon were consistently longer than unmarked Chinook salmon especially by June, but unmarked salmon had significantly greater body condition (based on length-weight residuals) for over half of the May sampling efforts. Both unmarked and marked fish ate similar types and amounts of prey for small (station) and large (month, year) scale comparisons, and feeding intensity and growth were not significantly different between the two groups. There were synchronous interannual fluctuations in catch, length, body condition, feeding intensity, and growth between unmarked and hatchery fish, suggesting that both groups were responding similarly to ocean conditions.     

Juvenile growth and aggression in diploid and triploid Chinook salmon Oncorhynchus tshawytscha (Walbaum)

Multilocus heterozygosity, aggressive and feeding behaviour, plasma cortisol levels and growth rate were evaluated among three groups of juvenile Chinook salmon Oncorhynchus tshawytscha : diploid, triploid and mixed groups of diploid and triploid fish. There was no difference between diploid and triploid fish in measurements of heterozygosity calculated using seven microsatellite loci, and these measurements did not correlate with performance measurements including feeding rate and growth rate. Aggression trials that examined small groups of fish revealed that after 4 days together in tanks, triploid fish were significantly less aggressive during feeding than diploid fish or fish in mixed groups. At the end of the trials, however, plasma cortisol levels did not differ among the three groups. Thirty-day growth trials in duplicate tanks of 60 fish revealed no difference in growth rate among diploid, triploid and mixed groups, but plasma cortisol levels were significantly lower in triploid fish than in either diploid fish or the mixed fish. Overall, independent of the above differences in aggressive behaviour and cortisol levels, these results suggest similar performance in diploid and triploid Chinook salmon, and thus provide support for the viability of triploid Chinook salmon culture in commercial aquaculture.     

Fast-growing transgenic common carp mounting compensatory growth

Compensatory growth is a phase of accelerated growth apparent when favourable conditions are restored after a period of growth depression. To investigate if F₂ common 'all-fish' growth hormone gene transgenic common carp ( Cyprinus carpio ) could mount compensatory growth, a 9 week study at 29° C was performed. The control group was fed to satiation twice a day throughout the experiment. The other two groups were deprived of feed for 1 or 2 weeks, respectively, and then fed to satiation during the re-feeding period. At the end of the experiment, the live masses of fish in the deprived groups were still significantly lower than those of the controls. During the re-feeding period, size-adjusted mean specific growth rates and mean feed intakes were significantly higher in the deprived fish than in the controls, indicating a partial compensatory growth response in these fish. No significant differences were found in food conversion efficiency between the deprived and control fish during re-feeding, suggesting that hyperphagia was the mechanism responsible for increased growth rates. The proximate composition of the deprived fish at the end of the experiment was similar to that of the control fish. This study is, to our knowledge, the first to report that fast-growing transgenic fish can achieve partial compensation of growth following starvation.     

Replacement of fish oil with thraustochytrid Schizochytrium sp. L oil in Atlantic salmon parr (Salmo salar L) diets

Replacing fish oil with that from a docosahexaenoic acid (22:6omega3, DHA) rich single cell micro-organism, thraustochytrid Schizochytrium sp. L, in diets for Atlantic salmon (Salmo salar) was investigated. Four experimental diets containing 100% thraustochytrid oil (TO), 100% palm oil (PO) and a 4:1 palm and thraustochytrid oil mixture (MX) were compared to a fish oil (FO) diet over 9 weeks. A saltwater transfer challenge occurred at the end of the trial for 14 days to test the diet treatments on the ability of salmon to smolt. There were no significant differences in the feed consumption of the diets or the digestibility of the omega3 or omega6 PUFA, indicating no differences in the digestibility of fatty acids between diets. No significant differences were noted between the growth of fish on the four diet treatments. Significant differences were noted in the fatty acid profiles of the fish muscle tissues between all diets. Fish on the TO diet had a significantly greater percentage of DHA in muscle tissue compared with fish on all other diets. Blood osmolarity, which is inversely related to the ability of salmon to smolt, from the TO and FO fed fish was significantly lower than that of fish on the PO diet. This study showed that thraustochytrid oil can be used to replace fish oil in Atlantic salmon diets without detriment to the growth of parr. Including thraustochytrid oil in fish diets significantly increases the amount of DHA in Atlantic salmon muscle and therefore is a candidate for use in oil blends for salmon diets. Thraustochytrid oil provides a renewable source of essential fatty acids, in particular DHA, for aquafeeds.     

Growth,survival, and tag retention of steelhead trout (<Emphasis Type="Italic">O. mykiss</Emphasis>) surgically implanted with dummy acoustic tags

Recent advances in micro-electronics make the study of the migration of even small marine animals (>12 cm) over many 1000s of kilometres a serious possibility. Important assumptions in long-term studies are that rates of tag loss caused by mortality or tag shedding are low, and that the tagging procedure does not have an unacceptable negative effect on the animal. This paper reports results from a study to examine the retention of relatively large (24 × 8 mm) surgically-implanted dummy acoustic tags over a 7-month period in steelhead pre-smolts (O. mykiss), and the effects of implantation on growth and survival. Although there was some influence on growth to week 12, survival was high for animals > 13 cm FL. In the following 16-week period, growth of surgically implanted pre-smolts was the same as the control population and there was little tag loss from mortality or shedding. Currently available acoustic tags can be implanted in salmonid fish ≥12 cm FL, although combined losses from mortality and tag shedding were 33–40% for animals in the 12 and 13 cm FL size classes. By 14 cm FL, combined rates of tag loss (mortality plus tag shedding) for surgically implanted tags dropped to <15% and growth following surgery was close to that of the controls. Our results suggest that studies of ocean migration and survival over periods of many months are now feasible even for animals as small as salmon smolts. Surgically implanted salmon smolts are therefore good candidates for freshwater and coastal ocean-tracking studies on relatively long time scales (months). On such time scales, even relatively small salmon smolts may move thousands of kilometers in the ocean.     

Otolith and somatic growth rates in Atlantic salmon parr, Salmo salar L: evidence against coupling

It is often assumed that otolith growth is in some way dependent on somatic growth (i.e. that the two processes are coupled). We examined the relationships between sagitta radius and fork length in 0+ Atlantic salmon parr that would subsequently smolt aged 1 + (UMG fish) or 2+ (LMG fish). Repeated measurements of fork lengths of individually marked parr, taken over a 211-day period from first feeding, were compared to sagitta radii on the same measuring dates (obtained by analysis of daily increments). The results showed that there was a linear relationship between fork length and otolith radius in UMG parr. However, this was not true for LMG parr. These fish enter a state of natural anorexia in their first autumn (despite excess food), but their otoliths continued to grow at the same rate despite the virtual cessation of somatic growth; they had therefore developed disproportionately large otoliths by the end of the study period. The relative growth rates of soma and otoliths first changed in LMG fish in late July/early August; this is the most precise estimate yet obtained of the timing of divergence in the developmental pathways of UMG and LMG parr. The rate of sagitta accretion was consistently lower in LMG parr, possibly indicating a lower metabolic rate in these fish. The results are discussed in relation to previous theories of the relationship between otolith and somatic growth.     

Plasticity of smolting in spring chinook salmon: relation to growth and insulin-like growth factor-I

In two year classes of Willamette River spring chinook salmon, reared at the Willamette Hatchery, and two groups of Yakima River spring chinook salmon, one sampled from the Yakima River and the other reared in a hatchery, fish which had relatively high growth rates in the summer–autumn period smolted in the autumn as measured by increases in gill Na⁺ K⁺ AT Pase activity. In contrast, groups with relatively low growth rate did not smolt in the autumn. Plasma levels of insulin-like growth factor-I (IGF-I) showed discrete differences between groups, with high levels associated with increased gill Na⁺ K⁺ AT Pase activities. These results demonstrate that smolting is plastic in spring chinook salmon, occurring in the autumn or the spring. In addition, smolting appeared to be related to growth rate; however, the relationships shown were correlational and causal mechanisms were not elucidated. Yet, the results do indicate a relationship between growth, an endocrine growth factor and smolting, suggesting a mechanistic link between developmental plasticity and the environment mediated by the endocrine system.     

Redistribution of juvenile salmonid fishes after localized catastrophic depletion

Juvenile Atlantic salmon and brown trout were depleted at three sites ( c . 108–380 m²) of a natural stream during the summer months of 1991 and 1992. Local population changes and movements of fish marked in sections adjacent to each depleted area were monitored thereafter. There was very little movement of marked salmon parr into the central regions of the depleted areas following the immediate post-marking period. Upstream movement by young-of-the-year fish from high density sections in mid-late summer was noted for trout but not salmon. Unmarked 1-year-old salmon parr immigrated into depleted areas in June 1992, and the pattern of recolonization was consistent with migration upstream from the adjoining river. It is concluded that resident salmon were very strongly site-attached and resource tracking was of no functional significance as a compensatory mortality mechanism. The occurrence of a long distance migratory component in the population during early-mid summer indicates that this, rather than local resource tracking, constitutes a potential compensatory mechanism.     

Effects of location of food delivery and social status on foraging-site selection by juvenile Atlantic salmon

Synopsis Experiments were conducted to investigate the response of juvenile Atlantic salmon,Salmo salar, to changes in the location at which food entered a stream tank. Experience with the location of food input into the system significantly influenced foraging-site selection by the dominant fish. Dominant fish changed location of their foraging site in response to a change in the location of food input, and occupied, aggressively defending, sites just downstream of the location of food introduction. In contrast, subordinate fish occupied foraging sites at the downstream end of the stream tank, regardless of location of food input. As a result of both site selection and social status, dominant fish captured significantly more prey than subordinates. Our results support the contention that salmonids choose foraging sites to maximize foraging opportunities. Our results also provide a possible explanation for the use of atypical foraging sites by individual fish within their home range over the course of a single day, as observed in a number of salmonid species in the field.