Maladaptation and phenotypic mismatch in cultured Atlantic salmon used for stocking

Compared to wild populations, cultured Atlantic salmon often sustain higher mortality rates and lower adult return rates when stocked as juveniles into natural streams. The ultimate causes for such differences in fitness, however, are not clear. Here we tested if relaxed levels of natural selection and improved survival in the hatchery environment could account for the observed degree of maladaptation among stocked fish. To do this, we assessed the degree of phenotypic mismatch between wild and cultured fish in three populations over five consecutive years. Significant differences were found in several phenotypic traits that are likely to have fitness implications. Thus, if the objective is to mimic wild individuals for restoration purposes, current hatchery practices aimed at maximising juvenile survival and enhancing growth may need to be revised.     

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.     

Morphological differences in parr of Atlantic salmon Salmo salar from three regions in Norway

Morphological characters were compared in parr (total length 33-166 mm) of Atlantic salmon Salmo salar sampled from eight wild populations in three regions, three in northern, two in the middle and three in southern Norway, covering a distance of 1700 km (from 70° N to 58° N). On the basis of morphological characters 94·6% of the individuals were correctly classified into the three regions. Discrimination between populations within these three regions also had a high degree of correct classification (89·0-95·8%). Principle component analysis identified largest differences to be in head characters, notably eye diameter and jawbone, with the smallest diameter and head size among the northernmost populations. Fish from the southern rivers had a deeper body form whereas fish from the middle region had larger heads and pectoral fins. This illustrates that S. salar already in the early parr stage has morphological traits, which can be used in discrimination between regions and populations and that these differences are discernible in spite of the volume of escaped farmed fish spawning in Norwegian rivers during the past 30 years.     

Sperm traits in farmed and wild Atlantic salmon Salmo salar

Differences in sperm metabolism and morphology between wild and non‐local farmed Atlantic salmon Salmo salar were assessed by measuring metabolic enzyme activities and length of sperm flagella. No differences were observed between wild and farmed S. salar sperm with regards to cell counts or any of the biochemical variables assessed. Flagella of sperm cells were significantly longer in wild than farmed S. salar; however, this did not result in higher energy levels or different fertilization rates.     

Multiple central-place territories in wild young-of-the-year Atlantic salmon Salmo salar

1. The space use of central-place foragers, animals that forage from and return to a single central place such as a nest, burrow or sleeping site, is well documented. Limited data, however, exist for multiple central-place foragers that alternate among several central places. 2. The conventional view of stream-dwelling salmonids suggests that they conform to the central-place territorial model (CPTM) by (i) attacking prey and intruders from one primary foraging station, and defending (ii) small (iii) exclusive areas that (iv) increase with body size. 3. Recent studies suggest greater variability in salmonid space-use than would be expected by the CPTM, but tend to focus on the time allocated towards different activities rather than their distribution in space, especially for young-of-the-year (YOY) fish that are hard to tag and monitor in the wild. 4. In this study, the validity of CPTM was tested by mapping the daily space use of 50 YOY Atlantic salmon in a natural stream via repeated observations of tagged fish in diverse habitats, and by comparing these to earlier estimates of territory use in YOY salmonids. 5. The 50 YOY Atlantic salmon were multiple central-place foragers. All fish visited more than one foraging station (median = 12.5 stations), visited most stations (68.5%) repeatedly, showed limited fidelity to a particular station and typically attacked prey only while holding a position at a station. 6. The multiple central-place territories of the 50 fish were large (mean = 0.932 m(2)) compared to earlier territory size estimates (mean = 0.107 m(2)) for salmonids of similar size and, surprisingly, did not increase with body size. Focal fish attacked intruders from similar distances as reported earlier for much smaller territories, suggesting that large territories are less exclusively defended at any given time. 7. Overall, this study provides a new view on foraging and territoriality in stream salmonids, and on the small but diverse literature on multiple central-place foragers. Further studies, however, are needed to clarify the evolutionary benefits and population consequences of multiple central-place space-use in mobile animals.     

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.     

Migration of wild and hatchery reared smolts of Atlantic salmon, Salmo salar L., through lakes

In 1982 and 1983 descending wild Atlantic salmon smolts, Salmo salar L., were caught in a fish-trap at the mouth of the River Imsa. Together with hatchery-reared smolts of the River Imsa strain they were tagged and released at three different sites of the Imsa-Lutsi watercourse: in the Imsa River 1 km above the trap, and in two lakes, 3 and 11 km upstream of the trap. The recapture-rate in the fish-trap decreased with increased migration distance. The hatchery-reared smolts migrated downstream faster than wild smolts. Lake-released smolts were considerably delayed in their downstream migration compared to the river released smolts. The results are discussed in relation to environmental and physiological parameters.     

Divergent trends in life‐history traits between Atlantic salmon Salmo salar of wild and hatchery origin in the Baltic Sea

Four Atlantic salmon Salmo salar stocks in the Baltic Sea, varying in their breeding history, were studied for changes in life‐history traits over the years 1972–1995. Total length (L_T) at age of captured (L_TC) fish had increased throughout the study period, partly due to increased temperature and increased L_T at release, (L_TR) but also due to remaining cohort effects that could represent unaccounted environmental or genetic change. Simultaneously, maturation probabilities controlled for water temperature, L_TC and L_TR had increased in all stocks. The least change was observed in the River Tornionjoki S. salar that was subject only to supportive stockings originating from wild parents. These results suggest a long‐term divergence between semi‐natural and broodstock‐based S. salar stocks. Increased L_T at age explained advanced maturation only marginally, and it remains an open question to what extent the generally increased probabilities to mature at early age reflected underlying genetic changes.     

The morphological plastic response to water current velocity varies with age and sexual state in juvenile Atlantic salmon, Salmo salar

1. Salmonids, like many other fish species, exhibit morphological plasticity to variations in water current velocity. However, little is known about how this response varies with age and alternative sexual tactics that usually coexist in the same area. We therefore sampled immature 1- and 2-year-old and sexually mature Salmo salar parr to determine how the morphological response to slow and rapid water currents varies across these groups.
2. Both 1- and 2-year-old immature parr in rapid habitats can be distinguished from individuals in slow habitats using a combination of fin measurements. In contrast, body shape measurements were useful only to distinguish 2-year-old individuals in the different habitat types. We also showed that mature parr are notably robust, irrespective of habitat type. For these individuals, only their body length differed between slow and rapid water currents, being bigger in slow water currents.
3. Our results imply that fins are the first structures to respond to water current velocity, followed by changes in body shape as individuals grow bigger. The robust phenotype observed for mature parr is likely to pose extra limitations on movement due to an increase in drag forces, thus contributing to their smaller size in rapid water currents.     

Genetic variation in threshold reaction norms for alternative reproductive tactics in male Atlantic salmon, Salmo salar

Alternative reproductive tactics may be a product of adaptive phenotypic plasticity, such that discontinuous variation in life history depends on both the genotype and the environment. Phenotypes that fall below a genetically determined threshold adopt one tactic, while those exceeding the threshold adopt the alternative tactic. We report evidence of genetic variability in maturation thresholds for male Atlantic salmon (Salmo salar) that mature either as large (more than 1 kg) anadromous males or as small (10-150 g) parr. Using a common-garden experimental protocol, we find that the growth rate at which the sneaker parr phenotype is expressed differs among pure- and mixed-population crosses. Maturation thresholds of hybrids were intermediate to those of pure crosses, consistent with the hypothesis that the life-history switch points are heritable. Our work provides evidence, for a vertebrate, that thresholds for alternative reproductive tactics differ genetically among populations and can be modelled as discontinuous reaction norms for age and size at maturity.     

Genetic determination of the contribution of stocked and wild Atlantic salmon, Salmo salar L., to the angling fisheries in two Spanish rivers

From 1983 to 1986 the Rivers Ason and Nansa (northern Spain) were stocked with over 100 000 eyed ova year⁻¹ from a Scottish hatchery in an effort to increase angling catches of Atlantic salmon. An inherent genetic marker was used to estimate the contributions of stocked and wild fish from these year-classes to the 1988 rod-and-line fishery. Stocking with foreign ova yielded 0.9 fish per 10 000 ova in the R. Asón and zero fish in the R. Nansa. Natural egg deposition yielded 4.1 and 4.9 fish per 10000 ova, respectively. Reasons for the difference in performance between stocked and wild fish are examined.     

Genetic assessment of Atlantic salmon Salmo salar and sea trout Salmo trutta stocking in a Baltic Sea river

Microsatellite DNA variation was used to assess the outcome of stocking Atlantic salmon Salmo salar and migratory trout Salmo trutta in River Sävarå, N Sweden. No information on pre‐stocking genetic composition of S. salar and S. trutta in River Sävarå was available. In 2 year‐classes of S. salar smolt, microsatellite data indicated that post‐stocking genetic composition differed markedly (F_ST= 0·048) from the main donor strain, Byskeälven S. salar, and from other Gulf of Bothnia S. salar stocks (F_ST 0·047 and 0·132). The STRUCTURE programme failed to detect any substructuring within Sävarå salmon. It was concluded that only minor introgression estimated to a proportion of 0·11 (95% CI 0·07–0·16) has occurred in S. salar. Salmo trutta showed overall low differentiation among populations with maximum F_ST of 0·03 making analysis more cumbersome than in S. salar. Still, the SävaråS. trutta deviated significantly from potential donor populations, and STRUCTURE software supported that majority of trout in Sävarå formed a distinct genetic population. Admixture was more extensive in S. trutta and estimated to 0·17 (95% CI 0·10–0·25).     

The survival of semi‐wild, wild and hatchery‐reared Atlantic salmon smolts of the Simojoki River in the Baltic Sea

The recapture rate and survival of hatchery‐reared Atlantic salmon Salmo salar stocked as 1 year‐old parr (semi‐wild) with that of hatchery‐reared Atlantic salmon stocked as 2 year‐old smolts and wild smolts of Atlantic salmon in the northern Baltic Sea were compared. This was done through tagging experiments carried out in 1986–1988 and 1992. The recapture rate of the semi‐wild groups varied from 1·0 to 13·1%, being similar in 3 tagging years and lower in 1 year than that of the wild groups (1·7–17·0%). The recapture rate of the semi‐wild groups was similar (in 2 years) or higher (in 2 years) than that of the hatchery‐reared groups stocked as smolts (1·3–6·3%). The survival of semi‐wild smolts during the sea migration was as high as that of wild Atlantic salmon of an equal size and two to three times higher than hatchery‐reared Atlantic salmon stocked as smolts. The survival rate was positively associated with smolt size. The suitability of hatchery‐reared parr and smolts in the management of reduced Atlantic salmon stocks is compared.     

Reduction in antipredator response detected between first and second generations of endangered juvenile Atlantic salmon Salmo salar in a captive breeding and rearing programme

Behaviour trials determining antipredator response were conducted on first and second generation juveniles from a captive breeding and rearing programme for endangered Inner Bay of Fundy Atlantic salmon Salmo salar. Second generation captive fry displayed significantly higher levels of risk‐taking behaviour before and after exposure to a simulated avian predator. Because the first and second generation fry were reared under the same environmental conditions and differed only in the number of generations spent in captivity, these results suggest that rapid genetic changes, possibly due to domestication selection, may have occurred. Antipredator response was also assessed in fully wild and highly domesticated experimental groups: wild fry displayed the greatest antipredator response and domesticated fry displayed the highest levels of risk‐taking behaviour. These results add to the growing evidence documenting rapid genetic change in response to rearing in a captive environment.     

Trans‐generational maternal effect: temperature influences egg size of the offspring in Atlantic salmon Salmo salar

Effect of increased temperature during egg maturation on the mass of single eggs produced by the offspring was investigated experimentally in Atlantic salmon Salmo salar. Mass of eggs produced by next‐generation females was larger when their mothers experienced warmer water during the last two months of egg maturation, relative to those that experienced unheated river water. There was no similar trans‐generational paternal effect on offspring egg mass.     

Passing a seawater challenge test is not indicative of hatchery‐reared Atlantic salmon Salmo salar smolts performing as well at sea as their naturally produced conspecifics

Despite satisfactory reactions to seawater challenge tests indicative of appropriate physiological state, hatchery‐reared Atlantic salmon Salmo salar smolts stocked in the Eira River in Norway between 2001 and 2011 performed less well at sea in terms of growth, age at maturity and survival than smolts of natural origin. The mean rates of return to the river for hatchery‐reared and naturally produced S. salar were 0·98 and 2·35%. In the Eira River, c. 50 000 hatchery‐reared S. salar smolts of local origin were stocked annually to compensate for reduced natural smolt production following regulation for hydroelectric purposes, while a mean of 17 262 smolts were produced naturally in the river. This study demonstrates that, although captive S. salar perform well in seawater challenge tests, hatchery‐reared smolts are not necessarily as adaptable to marine life as their naturally produced counterparts. These findings suggest that production of hatchery‐reared smolts more similar to naturally produced individuals in morphology, physiology and behaviour will be necessary to improve success of hatchery releases. Where possible, supplementary or alternative measures, including habitat restoration, could be implemented to ensure the long‐term viability of wild stocks.     

Radio‐telemetry shows differences in the behaviour of wild and hatchery‐reared European grayling Thymallus thymallus in response to environmental variables

Juvenile wild and hatchery‐reared European grayling Thymallus thymallus were tagged with radio‐transmitters and tracked in the Blanice River, River Elbe catchment, Czech Republic, to study their behavioural response to stocking and environmental variation. Both wild and hatchery‐reared T. thymallus increased their diel movements and home range with increasing light intensity, flow, temperature and turbidity, but the characteristics of their responses differed. Environmental variables influenced the movement of wild T. thymallus up to a specific threshold, whereas no such threshold was observed in hatchery‐reared T. thymallus. Hatchery‐reared fish displayed greater total migration distance over the study period (total migration) than did wild fish, which was caused mainly by their dispersal in the downstream direction.