Atlantic salmon Salmo salar L., brown trout Salmo trutta L. and Arctic charr Salvelinus alpinus (L.): a review of aspects of their life histories

Abstract – Among the species in the family Salmonidae, those represented by the genera Salmo, Salvelinus, and Oncorhynchus (subfamily Salmoninae) are the most studied. Here, various aspects of phenotypic and life‐history variation of Atlantic salmon Salmo salar L., brown trout Salmo trutta L., and Arctic charr Salvelinus alpinus (L.) are reviewed. While many strategies and tactics are commonly used by these species, there are also differences in their ecology and population dynamics that result in a variety of interesting and diverse topics that are challenging for future research. Atlantic salmon display considerable phenotypic plasticity and variability in life‐history characters ranging from fully freshwater resident forms, where females can mature at approximately 10 cm in length, to anadromous populations characterised by 3–5 sea‐winter (5SW) salmon. Even within simple 1SW populations, 20 or more spawning life‐history types can be identified. Juveniles in freshwater can use both fluvial and lacustrine habitats for rearing, and while most smolts migrate to sea during the spring, fall migrations occur in some populations. At sea, some salmon undertake extensive oceanic migrations while other populations stay within the geographical confines of areas such as the Baltic Sea. At the other extreme are those that reside in estuaries and return to freshwater to spawn after spending only a few months at sea. The review of information on the diversity of life‐history forms is related to conservation aspects associated with Atlantic salmon populations and current trends in abundance and survival. Brown trout is indigenous to Europe, North Africa and western Asia, but was introduced into at least 24 countries outside Europe and now has a world‐wide distribution. It exploits both fresh and salt waters for feeding and spawning (brackish), and populations are often partially migratory. One part of the population leaves and feeds elsewhere, while another part stays as residents. In large, complex systems, the species is polymorphic with different size morphs in the various parts of the habitat. Brown trout feed close to the surface and near shore, but large individuals may move far offshore. The species exhibits ontogenetic niche shifts partly related to size and partly to developmental rate. They switch when the amount of surplus energy available for growth becomes small with fast growers being younger and smaller fish than slow growers. Brown trout is an opportunistic carnivore, but individuals specialise at least temporarily on particular food items; insect larvae are important for the young in streams, while littoral epibenthos in lakes and fish are most important for large trout. The sexes differ in resource use and size. Females are more inclined than males to become migratory and feed in pelagic waters. Males exploit running water, near‐shore and surface waters more than females. Therefore, females feed more on zooplankton and exhibit a more uniform phenotype than males. The Arctic charr is the northernmost freshwater fish on earth, with a circumpolar distribution in the Holarctic that matches the last glaciation. Recent mtDNA studies indicate that there are five phylogeographic lineages (Atlantic, Arctic, Bering, Siberian and Acadian) that may be of Pleistocene origin. Phenotypic expression and ecology are more variable in charr than in most fish. Weights at maturation range from 3 g to 12 kg. Population differences in morphology and coloration are large and can have some genetic basis. Charr live in streams, at sea and in all habitats of oligotrophic lakes, including very deep areas. Ontogenetic habitat shifts between lacustrine habitats are common. The charr feed on all major prey types of streams, lakes and near‐shore marine habitats, but has high niche flexibility in competition. Cannibalism is expressed in several cases, and can be important for developing and maintaining bimodal size distributions. Anadromy is found in the northern part of its range and involves about 40, but sometimes more days in the sea. All charr overwinter in freshwater. Partial migration is common, but the degree of anadromy varies greatly among populations. The food at sea includes zooplankton and pelagic fish, but also epibenthos. Polymorphism and sympatric morphs are much studied. As a prominent fish of glaciated lakes, charr is an important species for studying ecological speciation by the combination of field studies and experiments, particularly in the fields of morphometric heterochrony and comparative behaviour.     

Resource partitioning between lake-dwelling Atlantic salmon (Salmo salar L.) parr, brown trout (Salmo trutta L.) and Arctic charr (Salvelinus alpinus (L.))

Abstract – Resource partitioning between Atlantic salmon parr, brown trout and Arctic charr was studied throughout the ice-free season in a north Norwegian lake. Juvenile salmon and trout (≤160 mm) utilized the littoral zone and juvenile charr the profundal, while adult trout and charr (>160 mm) were found in both. Juvenile salmon and trout had a similar diet, although trichopteran larvae were more important for the trout and chironomid pupae and three-spined sticklebacks for the salmon parr. Small salmon and trout parr (≤120 mm) had a higher diet overlap than larger parr (121–160 mm). The feeding habits of adult trout were similar to that of juvenile trout, but the former took larger prey items. At the population level, both salmon and trout were generalistic feeders with a broad diet, but at the individual level, both species had specialized on a single or a few prey categories. Juvenile charr were segregated from salmon and trout in both habitat and food utilization; they had a narrow diet consisting of chironomids and zooplankton, possibly reflecting their confinement to the profundal habitat which have a low diversity of potential prey. Larger charr also took zoobenthos and sticklebacks in the littoral zone. ^Note     

An estimation of dietary iron requirement of Atlantic salmon, Salmo salar L., parr

A study was conducted to determine the dietary iron requirement of fingerling Atlantic salmon Salmo salar L. During the first 4 weeks of the experiment, fish with an initial weight of 5 g were fed a casein–gelatine-based purified diet which contained 11 mg iron kg^?1. Thereafter duplicate tanks (200 fish in each) were fed the casein–gelatine purified diets containing supplemental iron levels of 0, 10, 20, 30, 40, 60, 100, 200 or 400 mg iron kg^?1 (added as FeSO_?4* 7H₂O) for 12 weeks. Weight gain, body length and mortality were monitored. Liver iron and ascorbic acid concentration were analysed in addition to whole-body iron, manganese and zinc concentration. Several haematological parameters were also measured. There were no significant differences in weight gain and survival of salmon fed diets containing different iron levels. Haematological values, hepatic and whole-body iron concentrations were, however, significantly affected by the dietary iron content. Liver vitamin C concentration decreased with increasing dietary iron levels. Dietary supplementation with iron significantly reduced whole-body manganese, but no effect of dietary iron on whole-body zinc was found. Based on haematology and hepatic iron concentration, the iron requirement of Atlantic salmon was determined to be between 60 and 100 mg iron kg¹.     

Control of bacterial kidney disease in Atlantic salmon, Salmo salar L., by dietary modification

Abstract. The effects of dietary treatments were investigated as a prophylactic measure to minimize the occurrence and severity of bacterial kidney disease (BKD) infections in Atlantic salmon, Salmo salar L., at Margaree Fish Culture Station, Nova Scotia, Canada. Six diets containing various levels of calcium, magnesium, zinc, iron, copper, manganese, cobalt and iodine in addition to one commercial diet were fed to post–yearling Atlantic salmon in two consecutive experiments. Natural infection was utilized to examine the effects of each diet on the prevalence of BKD. Diet containing high levels of iodine (4.5 mg/kg feed) and fluorine (4.5 mg/kg) reduced BKD prevalence rate to 3% and 5% respectively compared to 95% and 38% with commercial feed. Some other experimental diets also reduced prevalence of BKD, but to a lesser degree than the diet with additional iodine and fluorine.     

Influence of dietary vitamin B6 on tissue vitamin B6 contents and immunity in Atlantic salmon, Salmo salar L.

Atlantic salmon, Salmo salar L., (14 g) were fed a practical fish-meal-based diet supplemented with 0. 10, 20. 40, 80 and 160 mg pyridoxine (PN) per kg feed for 20 weeks. Tissue vitamin B₆ contents were significantly reduced in fish fed the non-supplemented diet compared with fish fed PN-supplemented diets. Serum haemolytic complement activity and head kidney lysozyme activity, and the specific antibody response following immunization with Vibrio salmonicida. were not influenced by the dietary regimes. Challenge with Aeromonas salmonicida showed that increasing the dietary levels of vitamin B₆ did not improve the resistance to furunculosis. Growth, mortality and haematology were not affected by supplementing a practical diet with vitamin B₆ In conclusion, feed levels of vitamin B₆ higher than the minimum dietary requirement did not enhance immune functions and disease resistance in Atlantic salmon.     

Atlantic salmon, Salmo salar L., feed in Scottish coastal waters

Abstract. The stomach contents of 256 Atlantic salmon, Salmo salar L., of fork lengths 53–66cm, caught in bag nets on the Scottish west coast near Ullapool, were examined between June 1983 and July 1986. A total of 61 fish contained fish in the stomach or had faecal pellets containing fish bones in the gut. All recognizable whole fish were sandeels, Ammodytes marinus Raitt, ranging in size from 4.5 to 15cm. No evidence for crustacean or other non-fish prey items was found. Results indicate that feeding salmon were caught up to a certain cut-off point in June or early July, after which all salmon sampled were not feeding. It is suggested that either there is a local feeding stock of salmon or that fish feed during migration from the Faroe Isles or other possible distant water origins.     

Screamer disease in Atlantic salmon, Salmo salar L., in Chile

Investigation of cranial and spinal deformities in Atlantic salmon smolts from Chile immediately after sea transfer, over two successive crops, demonstrated ankylosis of the mandibular articulation, spinal foreshortening, fracture of vertebrae and rarefaction of osseous and cartilaginous tissues including the operculum. As a result the mouth was permanently agape and opercula could be folded back on themselves. All affected fish had been transferred to sea water at 20 °C plus and fed on particular commercial diets. Fish in cooler areas, or on diets high in vitamin C and phosphorus in the high temperature zone, were clinically normal. The condition has not recurred following dietary adjustment.     

Pathogenicity of Saprolegnia spp. to Atlantic salmon, Salmo salar L., eggs

Live and dead Atlantic salmon eyed eggs were challenged with eight different Saprolegnia isolates, selected because of their varied origins, known morphological characteristics and growth/germination pattern. Some isolates were also tested for pathogenicity to Atlantic salmon parr. Challenge of eggs was performed by exposure to spores in suspension or by co-incubation of live eggs with infected dead eggs. The phenotypic characteristics of the isolates were evaluated in relation to their observed pathogenicity from the challenge experiment, to identify possible virulence factors leading to egg-infection by Saprolegnia. The results from the experiments confirm that live eggs are refractory to infection with Saprolegnia spores in suspension and that an infection of live eggs can only occur from an infection nucleus represented by dead eggs or debris. It was observed that strains pathogenic to salmon parr were not particularly infective towards eggs, and the isolates that gave the highest infection rates to eggs were species considered to be saprotrophs.     

Comparative virulence of Infectious salmon anaemia virus isolates in Atlantic salmon, Salmo salar L.

Infectious salmon anaemia virus (ISAV) surveillance in the Bay of Fundy has identified the existence of a large number of genetically distinct ISAV isolates which appear to be of variable virulence. Genetically distinct isolates are currently being designated based on sequencing of the hyper polymorphic region (HPR) of genomic segment 6, which encodes the haemagglutinin–esterase protein, but it has been difficult to elucidate a clear association between these molecular variations and variations in virulence. This has hampered the establishment of proactive management decisions regarding infected fish, and ISAV infections, regardless of type, must be treated as one. Field data of ISAV infections is difficult to collect and to compare between infections because of a wide range of confounding factors including time of year, fish stock, cage site location, mitigating factors and stressors. An important tool in determining the relationship between molecular differences and virulence comes from analysis of quarantine studies. The goal of this study was to compare the virulence, by co-habitation and intraperitoneal injection, of four regionally common and recent ISAV isolates in a controlled environment. We found significant differences in mortality between ISAV molecular isolates, and present data showing that survival of ISAV infection confers significant resistance to re-infection with a different ISAV isolate. These findings, if borne out in field studies, will significantly alter the way ISAV infections are managed in the Bay of Fundy and elsewhere.     

Nocardiosis in tank-reared Atlantic salmon, Salmo salar L.

Nocardiosis is an infection caused by aerobic Gram-positive, branching, filamentous rods of the genus Nocardia . The organism is resident in both soil and plants ( Austin & Austin 1993 ; Frerichs 1993 ), and is closely related to Mycobacterium spp. Nocardiosis caused by Nocardia asteroides and N. seriolae (previously N. kampachi ) has been reported in several finfish species, both freshwater and marine. The first reported incidence was recorded by Valdez & Conroy (1963) in neon tetras, Hyphessobrycon innesi (Myers), and subsequently in other fish species, some of commercial significance including rainbow trout, Oncorhynchus mykiss (Walbaum) ( Snieszko, Bullock, Dunbar & Pettijohn 1964 ), brook trout, Salvelinus fontinalis (Mitchill), ( Campbell & MacKelvie 1968 ) and yellowtail, Seriola quinqueradiata (Temminck & Schlegel; Kubota, Kariya, Nakamura & Kira 1968 ). Despite two of these incidences occurring in salmonids, infection by Nocardia spp. in Atlantic salmon, Salmo salar L., has not previously been reported.     

Reproductive strategies in Atlantic salmon, Salmo salar L.

Abstract. Life-history strategies are means by which animals solve the problems of successful reproduction in varying environments. Their development patterns are consequences of responses to the opportunities the environment offers them. Understanding them requires an understanding of the way they evolved, their ontogenetic development, their physiological control, and their adaptive value. The present paper views the salmonids as marine fishes, which have radiated into fresh water through using river beds as protected spawning grounds. It also takes the view that the maturation process has priority over somatic growth in fish, and that it has already been initiated by the time of first feeding. Its completion is environmentally dependent, and can be arrested annually. Whether or not it will be arrested depends on the status of the energy stores of the individual at particular critical times of year. This mechanism has adaptive value both for immediate reproductive success — adequate energy to provision the next generation — and for later overwinter survival, ensuring that if energy stores are inadequate for reproduction they are spared. Atlantic salmon show variation in their reproductive patterns, and examples are given from laboratory and aquaculture experiments to demonstrate some environmental controls which result in these variations. A hypothetical model is presented to account for the operation of these controls.     

Potential climate change impacts on Arctic char Salvelinus alpinus L. in Ireland

Climate change has been identified as a global threat to Arctic char Salvelinus alpinus L. populations. Bayesian statistical models were used to identify important physical and ecological factors explaining Arctic char presence and persistence in Irish lakes. Maximum lake depth and the presence of mixed fish communities (i.e. including non‐salmonid fish species) were the most important drivers of Arctic char presence. There was a 75% probability that an Irish lake would contain Arctic char when maximum depth exceeded 40 m, if only a salmonid community was present. However, the required depth increased to 57 m when a mixed fish community (e.g. pike, perch or roach) was present. Similar variables explained char persistence, with surviving Arctic char populations being more likely in lakes with greater maximum depth and fewer non‐salmonid fish species. Tested temperature covariates were not important in explaining Arctic char presence or persistence in Irish lakes. A risk analysis was undertaken to identify which Arctic char lakes are vulnerable to colonisation by thermally plastic mixed fish communities due to flooding. Results indicated that 32 out of 45 lakes were not at any risk from the colonisation of thermally plastic mixed fish communities and 13 lakes had some level of risk.     

Habitat utilization of juvenile Atlantic salmon (Salmo salar L.), brown trout (Salmo trutta L.) and Arctic charr (Salvelinus alpinus (L.)) in two lakes in northern Norway

Abstract— Habitat utilization of juvenile Atlantic salmon, brown trout and Arctic charr was investigated in two lakes in northern Norway during the icefree season. Both the vertical distribution and the distribution among different habitat types were studied by gillnetting with small mesh sized gillnets (8-15 mm) in different habitats. Salmon and trout were predominantly caught in the littoral and sublittoral zones (0-6 m depth). Access to shelter seemed to be the most important factor determining the horizontal distribution of small salmon and trout. Most of these fish were caught in stony or vegetated habitats, while few salmon and trout were caught on sandy locations or in the pelagic zone. In one of the lakes, there were significantly higher catch rates of salmon than of trout in the stony littoral (0-3 m), while in the other lake there were no significant differences in spatial distribution between these two species. Charr were primarily found in the profundal, sublittoral or pelagic zones of the lakes.     

The effects of siltation on Atlantic salmon, Salmo salar L., embryos in the River Bush

The tolerance of incubating salmon embryos to spawning gravel sedimentation was examined under hatchery conditions and also in the natural state on the River Bush, County Antrim, Northern Ireland. In a laboratory assessment alevin survival was closely related to the level of fine material. The number of individuals was reduced at the 10% fines level while >15% fine material was found to be deleterious to survival. Mean survival on the river (19.26%) was found to be better than that recorded in the hatchery, although no clear relationship was established between the level of fines and percentage survival in the wild. The mean fines accumulation (13.59%) in the river incubators at the end of the experiment was shown to be statistically similar to the background substrate. The role of high flow events in the contamination of gravels scoured by spawning fish is discussed.     

Growth and survival of Atlantic salmon, Salmo salar L., fed different dietary levels of astaxanthin. First-feeding fry

Atlantic salmon fry hatched from pigment-free eggs and from eggs containing the pigment astaxanthin were fed eleven casein/gelatine-based purified diets with varying levels of astaxanthin, ranging from 0 to 317 mg kg^?1, to determine the optimum dietary astaxanthin level for satisfactory growth and survival during the start-feeding period. The fish were fed the experimental diets for a period of 11 weeks. No difference in performance was found between the two types of fry originating from the pigment-free eggs and those containing pigment. However, the dietary astaxanthin concentration was found to have a significant effect on both the growth and the survival of fry. Fish fed diets with astaxanthin concentrations below 5.3 mg kg^?1 were found to have marginal growth. In addition, mortality was high in the groups fed diets with astaxanthin concentrations below 1.0 mg kg^?1. The specific growth rate (SGR) was also affected by the dietary treatment. The lipid content was higher and the moisture content was lower in the fish fed the diets containing astaxanthin concentrations above 5.3 mg kg^?1. The vitamin A and astaxanthin concentrations in whole-body samples of the fry were significantly affected by the dietary level of astaxanthin. A plateau level in whole-body vitamin A concentration was observed at dietary levels of approximately 80 mg astaxanthin kg^?1 and higher, while no maximum astaxanthin concentration in whole-body samples was observed within the dietary levels used. The results suggest the need for a minimum dietary astaxanthin concentration of 5.1 mg kg^?1 to achieve maximum growth and survival during the start-feeding period. The results indicate a low bioavailability of vitamin A palmitate and acetate and the results also suggest a provitamin A function for astaxanthin during the same period.     

Glucose tolerance in Atlantic salmon, Salmo salar L., dependence on adaption to dietary starch and water temperature

Individually tagged Atlantic salmon, Salmo salar L., were fed three different adaptation diets containing low, medium or high levels of extruded starch, before intraperitoneal injection of glucose. The injections were done at winter and at summer temperatures. This was to investigate whether the tolerance for glucose varied according to adaption diet and/or according to season. The results showed a temperature-dependent response. At winter temperatures (2°C) the Atlantic salmon had problems adapting to both medium and high levels of dietary starch. This was indicated by lower growth response in the adaptation period, as well as by delayed glucose regulation. At summer temperatures (12.5°C) the Atlantic salmon could tolerate and utilize both medium and high levels of dietary starch. In summer also all groups responded evenly to the tolerance tests with respect to blood glucose regulation. Incorporation of liver glycogen was higher at summer than at winter temperatures in the high- carbohydrate group. No effect was found on haematocrit values as a response to diet or season.