Seasonal variation in the thermal performance of juvenile Atlantic salmon (Salmo salar)

1. Experimental data on the maximum growth and food consumption of winter‐acclimatised Atlantic salmon (Salmo salar) juveniles from three Norwegian rivers situated at 59 and 70°N were compared with predictions from published models of growth and food consumption of summer‐acclimatised fish from the same populations. 2. All winter‐acclimatised fish maintained positive growth and a substantial energy intake over the whole range of experimental temperature (1–6 °C). This contrasted with predictions from growth models based on summer acclimatised Atlantic salmon, where growth and energy intake ceased at approximately 5 °C. 3. Growth and food consumption varied significantly among populations. Winter‐acclimatised fish from a Northern population had a higher mass‐specific growth rate, higher energy intake and higher growth efficiency than southern populations, which is contrary to predictions from models developed using summer‐acclimatised salmon, where fish from the Northern population had the lowest growth efficiency. 4. The experiment provides evidence that thermal performance varies seasonally and suggests adaptation to the annual thermal regime.     

Territory size and distribution of newly-emerged Atlantic salmon (Salmo salar)

Newly emerged Atlantic salmon (Salmo salar) were observed from May to August 1981 at six isolated redds in Washington County, Maine, USA. Territorial size and distribution were measured. At the end of the emergence period (12 to 28 May), fish maintained positions (stations) at redds where water velocity did not exceed 52 cm s^–1 By 12 June, most salmon (80–96%) had moved off the redds of origin and had established territories 1 to 5 m from the redd. The area defended increased substantially after mid-June, but territorial aggression diminished by 15 July, and the fry dispersed downstream. All fish observed were territorial, and the percentage of time during which stations were held decreased from 89 in mid-May to as low as 40% in mid-June.Cooperators are the Maine Department of Inland Fisheries and Wildlife, University of Maine, U.S. Fish and Wildlife Service, and the Wildlife Management Institute     

Atlantic salmon (Salmo salar) of the Pigou and Bouleau Rivers,Quebec

Synopsis Juvenile Atlantic salmon of the Pigou and Bouleau Rivers, Quebec, were studied during the summer of 1972 and 1973. Growth increments for Pigou salmon averaged 36–50 mm yr^–1 and smolts averaged 2.9 yrs old. Bouleau salmon grew slower (26–41 mm yr^–1) and became smolts at a slightly older mean age (3.1 yrs). Mature male parr were more abundant in all age-groups from the Plgou River, possibly because of the faster growth. In terms of management, the Pigou River has little potential for increased salmon production because of limited habitat. On the other hand, the Bouleau River has considerable potential, and salmon production could be increased if the upper reaches were made accessible to migrating salmon.Contribution number 36 of the Matamek Research Station, Woods Hole Oceanographic Institute.     

Influence of size, winter duration and density on sexual maturation of Atlantic salmon (Salmo salar) juveniles in Little Codroy River (southwest Newfoundland)

Factors affecting sexual maturation of Atlantic salmon ( Salmo salar L.) juveniles in fresh water were analysed under wild conditions in Little Codroy River (southwest Newfoundland). Large size after the first year of life and short winters favoured maturation of age 1 + male parr. Incidence of maturing 1+ males was greater where high densities of parr (>0.2 individuals m ²; fish >6cm fork length) were recorded in the second season of growth. These results were discussed according to current knowledge on early maturation in Atlantic salmon.     

Selection against late emergence and small offspring in Atlantic salmon (Salmo salar)

Timing of breeding and offspring size are maternal traits that may influence offspring competitive ability, dispersal, foraging, and vulnerability to predation and climatic conditions. To quantify the extent to which these maternal traits may ultimately affect an organism's fitness, we undertook laboratory and field experiments with Atlantic salmon (Salmo salar). To control for confounding effects caused by correlated traits, manipulations of the timing of fertilization combined with intraclutch comparisons were used. In the wild, a total of 1462 juveniles were marked at emergence from gravel nests. Recapture rates suggest that up to 83.5% mortality occurred during the first four months after emergence from the gravel nests, with the majority (67.5%) occurring during the initial period ending 17 days after median emergence. Moreover, the mortality was selective during this initial period, resulting in a significant phenotypic shift toward an earlier date of and an increased length at emergence. However, no significant selection differentials were detected thereafter, indicating that the critical episode of selection had occurred at emergence. Furthermore, standardized selection gradients indicated that selection was more intense on date of than on body size at emergence. Timing of emergence had additional consequences in terms of juvenile body size. Late-emerging juveniles were smaller than early-emerging ones at subsequent samplings, both in the wild and in parallel experiments conducted in seminatural stream channels, and this may affect success at subsequent size-selective episodes, such as winter mortality and reproduction. Finally, our findings also suggest that egg size had fitness consequences independent of the effects of emergence time that directly affected body size at emergence and, in turn, survival and size at later life stages. The causality of the maternal effects observed in the present study supports the hypothesis that selection on juvenile traits may play an important role in the evolution of maternal traits in natural populations.     

Effects of sexual maturation, castration, and androgen implants on growth in one- and two-year-old parr in a Baltic Atlantic salmon (Salmo salar L.) stock

The effects of maturation, castration, and androgen implants on growth in 1- and 2-year-old Baltic salmon ( Salmo salar L.) were studied in a stock originating from the Umeälven (Ume river) in northern Sweden. 1 + male parr that later matured were larger than non-maturing parr in the spring. Later in the summer, during the period of gonadal growth, the growth rate of maturing parr was depressed compared to that of non-maturing males. Implantation of capsules with 11-ketoandrostenedione (OA) or testosterone (T) in sexually immature 1-year-old fish, accelerated the specific growth rate in spring and early summer, particularly considering growth in weight, resulting in an increase in the condition factor. In contrast, the growth rate was depressed from late June onwards, the period when the gonads normally develop in 1 + parr. T, but not OA, also promoted sexual maturation. Castration or OA treatment did not affect total summer growth in 2-year-old salmon. This study suggests a causal relationship between androgens and the growth pattern observed in maturing 1 + male parr.     

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.     

Catabolism of circulating collagen in the Atlantic salmon (Salmo salar)

The catabolic fate of circulating collagen (Col) in the Atlantic salmon (Salmo salar) was studied. Serum t_1/2 and organ distribution of circulating Col in salmon were determined using Col conjugated with ^l25I-tyramine cellobiose (¹²⁵I-TC), a low molecular weight adduct which is trapped intralysosomally at the site of uptake. Intravenously administered ^l25I-tyramine cellobioselabelled Col type I was prepared either from salmon skin (sCol) or rat skin (rCol). Biphasic clearance kinetics of ^l25I-TC-sCol in salmon were apparent, with 78% being removed from the circulation in an initial rapid α-phase (t_1/2(α) = 2.4 min), and 22% being removed more slowly in a terminalβ-phase (t]2(β) = 25.8 min). Serum half life of ¹²⁵I-TC-rCol was found to be 5.4 min (in this type of experiment the number of data points allow the determination of only a monophasic decay slope). Approximately 90% of recovered radioactivity was found in the kidney of the fish. In comparative experiments, 74% of administered ¹²⁵I-TC-sCol was cleared from the circulation of rats during an initial rapid α-phase with t_l/2(α) = 0.8 min, and 26% was eliminated in the terminal β-phase with t_1/2(β) = 7.2 min ^l25I-sCol was endocytosed and degraded in pure cultures of ral liver endothelial cells, which are the main site of clearance of circulating Col in the rat. Moreover, Col from the two species competed for the same receptor on cultured rat liver endothelial cells, Intravenous administration of tetramethyl rhodamine isothiocyanate-labelled sCol (TRITC-sCol) in salmon, and subsequent examination of sections of kidney in the fluorescence microscope, revealed that the fluorochrome was accumulated exclusively in discrete vesicles of sinusoidal lining cells. Analyses of kidney tissue 24h after intravenous administration of a mixture of fluorescein isothiocyanate-labelled latex beads and TRITC-sCol revealed no codistribution of the two fluorochromes, suggesting that the injected Col was taken up in cells different from macrophages. Purified pronephros macrophages prepared after simultaneous injections of stained beads and Col contained only fluorescein-labelled latex particles. Interestingly, the cells which had accumulated TRITC-sCol appeared to be equally distributed in both pronephros and the part of the kidney containing tubuli. We conclude that Col which gains access to the circulation of the Atlantic salmon is cleared mainly by uptake into sinusoidal lining cells of the kidney. These cells are distinct from phagocytosing macrophages, and morphologically similar to the highly specialized scavenging endothelial cells of mammalian liver sinusoids.     

Differences in growth between maturing and non-maturing male Atlantic salmon, Salmo salar L., parr

Sibling male Atlantic salmon parr that matured tended to be the larger fish in January, but their monthly specific growth rates between January and July did not differ from those of non-maturing fish. Maturing fish had lower condition factors in March, but greater increases in condition factor during April, exceeding those of non-maturing males by May. In maturing males, feeding rates between July and September, and specific growth rates in August and September, were lower than those of immature fish. Consequently, the mean size of immatures equalled or exceeded that of maturing males by October. Maturation rates were strongly correlated with increases in mean condition factor only during April.     

Atlantic salmon (Salmo salar L.) genetics in the 21st century: taking leaps forward in aquaculture and biological understanding

Atlantic salmon (Salmo salar L.) is among the most iconic and economically important fish species and was the first member of Salmonidae to have a high‐quality reference genome assembly published. Advances in genomics have become increasingly central to the genetic improvement of farmed Atlantic salmon as well as conservation of wild salmon stocks. The salmon genome has also been pivotal in shaping our understanding of the evolutionary and functional consequences arising from an ancestral whole‐genome duplication event characterising all Salmonidae members. Here, we provide a review of the current status of Atlantic salmon genetics and genomics, focussed on progress made from genome‐wide research aimed at improving aquaculture production and enhancing understanding of salmonid ecology, physiology and evolution. We present our views on the future direction of salmon genomics, including the role of emerging technologies (e.g. genome editing) in elucidating genetic features that underpin functional variation in traits of commercial and evolutionary importance.     

Links between patterns of marine growth and survival of Atlantic salmon Salmo salar, L.

The hypothesis that marine survival of Atlantic salmon Salmo Salar is linked to marine growth was explored by using inter-circuli distances and total numbers of circuli existing on scales from a population monitored over nearly four decades. The results suggest that marine growth controls survival, particularly during the late summer and early winter of the first year at sea. Recruitment is strongly linked to growth, described as the total number of circuli, but not to inter-circuli distances. This highlights the potential of patterns of circuli number to be considered as proxies for growth. Indications that hatchery populations might be subject to other mortality events, in addition to those experienced by wild populations, are also presented.     

Sexual maturity increases mobility and heterogeneity in individual space use in Atlantic salmon (Salmo salar) parr

We assessed the effects of sexual maturity on space use in Atlantic salmon (Salmo salar) parr as facultative early maturation enables us to work on individuals belonging to the same cohort. We monitored the space use of 40 1-year-old males in natura throughout a breeding season. First, mature individuals covered longer distances (absolute and upstream) and located within broader home ranges than immature parr. Second, sexual maturity also generated a higher interindividual variability in space use. Finally, mature individuals exhibited a higher probability of association with likely breeding sites on average. However, some mature individuals experienced a lower probability than immature individuals, suggesting that the space use of some mature individuals may not be optimal. Moreover, mature parr exploiting a broader home range or covering longer upstream distances had a higher probability of association with likely breeding sites. Covering longer upstream distances may therefore increase the reproductive success of mature parr, while involving higher energetic costs and a greater risk of predation.     

A linkage map of Atlantic salmon (Salmo salar) reveals an uncommonly large difference in recombination rate between the sexes

A genetic linkage map of the Atlantic salmon (Salmo salar) was constructed, using 54 microsatellites and 473 amplified fragment length polymorphism (AFLP) markers. The mapping population consisted of two full-sib families within one paternal half-sib family from the Norwegian breeding population. A mapping strategy was developed that facilitated the construction of separate male and female maps, while retaining all the information contributed by the dominant AFLP markers. By using this strategy, we were able to map a significant number of the AFLP markers for which all informative offspring had two heterozygous parents; these markers then served as bridges between the male and female maps. The female map spanned 901 cM and had 33 linkage groups, while the male spanned 103 cM and had 31 linkage groups. Twenty-five linkage groups were common between the two maps. The construction of the genetic map revealed a large difference in recombination rate between females and males. The ratio of female recombination rate vs. male recombination rate was 8.26, the highest ratio reported for any vertebrate. This map constitutes the first linkage map of Atlantic salmon, one of the most important aquaculture species worldwide.     

Bayesian genome‐wide association analysis for body weight in farmed Atlantic salmon (Salmo salar L.)

We performed a genome‐wide association study to detect markers associated with growth traits in Atlantic salmon. The analyzed traits included body weight at tagging (BWT) and body weight at 25 months (BW25M). Genotypes of 4662 animals were imputed from the 50K SNP chip to the 200K SNP chip using fimpute software. The markers were simultaneously modeled using Bayes C to identify genomic regions associated with the traits. We identified windows explaining a maximum of 3.71% and 3.61% of the genetic variance for BWT and BW25M respectively. We found potential candidate genes located within the top ten 1‐Mb windows for BWT and BW25M. For instance, the vitronectin (VTN) gene, which has been previously reported to be associated with cell growth, was found within one of the top ten 1‐Mb windows for BWT. In addition, the WNT1‐inducible‐signaling pathway protein 3, melanocortin 2 receptor accessory protein 2, myosin light chain kinase, transforming growth factor beta receptor type 3 and myosin light chain 1 genes, which have been reported to be associated with skeletal growth in humans, growth stimulation during the larval stage in zebrafish, body weight in pigs, feed conversion in chickens and growth rate of sheep skeletal muscle respectively, were found within some of the top ten 1‐Mb windows for BW25M. These results indicate that growth traits are most likely controlled by many variants with relatively small effects in Atlantic salmon. The genomic regions associated with the traits studied here may provide further insight into the functional regions underlying growth traits in this species.     

Migratory and reproductive behaviour of female adult Atlantic salmon, Salmo salar L., in a spawning stream

Migration and spawning behaviour of eight Atlantic salmon adult females were analysed by radio-tracking in relation to the degree of sexual maturity in a spawning tributary of the R. Sélune. Six of them were grilse and four of them were ripe. All the fish migrated upstream until reaching their spawning site at a distance of 4–12 km from the trap. The daily migration rate up to this site was inversely correlated with the length of the female. Spawning occurred in all fish at the same time when the water temperature increased dramatically. Spawning lasted 1–10 days according to the fish. After spawning, females quickly moved downstream only small distances and then stayed in approximately the same location until death. This study concluded that ripeness did not influence behaviour on the spawning migration and describes certain characteristics of the reproductive phase in a spawning tributary.     

A multi-population approach supports common patterns in marine growth and maturation decision in Atlantic salmon (Salmo salar L.) from southern Europe

This study provides a regional picture of long-term changes in Atlantic salmon growth at the southern edge of their distribution, using a multi-population approach spanning 49 years and five populations. We provide empirical evidence of salmon life history being influenced by a combination of common signals in the marine environment and population-specific signals. We identified an abrupt decline in growth from 1976 and a more recent decline after 2005. As these declines have also been recorded in northern European populations, our study significantly expands a pattern of declining marine growth to include southern European populations, thereby revealing a large-scale synchrony in marine growth patterns for almost five decades. Growth increments during their sea sojourn were characterized by distinct temporal dynamics. At a coarse temporal resolution, growth during the first winter at sea seemed to gradually improve over the study period. However, the analysis of finer seasonal growth patterns revealed ecological bottlenecks of salmon life histories at sea in time and space. Our study reinforces existing evidence of an impact of early marine growth on maturation decision, with small-sized individuals at the end of the first summer at sea being more likely to delay maturation. However, each population was characterized by a specific probabilistic maturation reaction norm, and a local component of growth at sea in which some populations have better growth in some years might further amplify differences in maturation rate. Differences between populations were smaller than those between sexes, suggesting that the sex-specific growth threshold for maturation is a well-conserved evolutionary phenomenon in salmon. Finally, our results illustrate that although most of the gain in length occurs during the first summer at sea, the temporal variability in body length at return is buffered against the decrease in post-smolt growth conditions. The intricate combination of growth over successive seasons, and its interplay with the maturation decision, could be regulating body length by maintaining diversity in early growth trajectories, life histories, and the composition of salmon populations.     

The relation between stock size and progeny of Atlantic salmon, Salmo salar L., in a Scottish stream

Large variations in the number of female adult salmon spawning in the Girnock Burn each year (range 28–127) produced smaller variations each year (range 2900–5600) in the number of juvenile migrants. The relative constancy of the parr migrations was achieved by changes in their age composition. There may be no advantage in allowing ova deposition to exceed a level around 200 000 (3.4 m⁻²) in the Girnock Burn, which would give rise to an average of about 4000 juvenile migrants (0.07 m⁻²) per season. Large numbers of ova (up to 12.5 m⁻²) did not decrease the number of juvenile migrants.