The influence of previous feeding regime on the compensatory growth response of maturing and immature Arctic charr, Salvelinus alpinus

Alternating periods of food deprivation with those of unlimited provision of food depressed the growth of Arctic charr, Salvelinus alpinus , below that of controls. Fish that were deprived of food and then fed on alternate weeks (1:1) were larger than those that were exposed to periods of 1 5- or 3-week deprivation and feeding (1·5:1·5 or 3:3). On receiving excess food supplies following 24 weeks on the restricted feeding regimes the previously-restricted fish grew more rapidly than the controls. The greatest compensatory growth was displayed after the 3:3 regime, followed by the 1·5: 1·5 and then the 1:1 feeding regime. At the termination of the experiment there were no significant differences in body weight between fish fed according to the different regimes during the period that food restriction was imposed. Growth patterns of the immature males and females were similar, but mature males were significantly lighter than the immature fish by the end of the experiment. Both immature and maturing fish displayed a compensatory growth response on return to adequate feeding. Beginning food restriction in May did not influence the proportions of male fish ( c . 60%) which were mature in the autumn.     

Effect of feeding frequency on food intake and growth of Arctic charr, Salvelinus alpinus L.

Social interactions and dominance hierarchy effects are important factors governing rates of growth of Arctic charr, Salvelinus alpinus L. The effects of hierarchy were increased as access to food became more restricted, i.e. feeding frequency was reduced, but these effects could not be attributed to direct competition for food since fish were fed to satiation at each feeding period. The results suggest that, whilst some fish on the restricted feeding regime were able to maintain good rates of growth, feeding by the majority of the fish was inhibited by the presence of larger individuals. Due to the importance of these hierarchy effects it was not possible to demonstrate physiological adaptations in fish allowed infrequent access to food.     

The effects of feeding regime on proximate body composition and patterns of energy deposition in juvenile Arctic charr, Salvelinus alpinus

Juvenile Arctic charr, Salvelinus alpinus , subjected to either food deprivation or restricted feeding showed reductions in relative sizes of both liver and viscera. With food deprivation, percentage lipid decreased and percentage water increased in the eviscerated carcass, which is in agreement with findings for several other fish species. Following transfer from a restricted to a satiation feeding regime, the relative sizes of both liver and viscera increased to levels exceeding those of fish fed continuously to satiation. At the end of the experiment the percentage liver lipid content of restricted-satiation fed fish was higher than in fish fed to satiation. These differences in energy deposition patterns were, however, not sufficiently large to produce marked differences at the whole animal level, since whole body lipid energy: protein energy ratios and energy utilization efficiencies were similar for restricted-satiation and satiation fed fish.     

Digestive capacities,inbreeding and growth capacities in juvenile Arctic charr Salvelinus alpinus

Genetic variation in growth performance was estimated in 26 families from two commercial strains of Arctic charr Salvelinus alpinus. Physiological determinants of growth and metabolic capacities were also assessed through enzymatic assays. A relatedness coefficient was attributed to each family using parental genotypes at seven microsatellite loci. After 15 months of growth, faster growing families had significantly lower relatedness coefficients than slower growing families, suggesting their value as indicators of growth potential. Individual fish that exhibited higher trypsin activity also displayed higher growth rate, suggesting that superior protein digestion capacities can be highly advantageous at early stages. Capacities to use amino acids as expressed by glutamate dehydrogenase (GDH) activities were lower in the liver of fast‐growing fish (13–20%), whereas white muscle of fast‐growing fish showed higher activities than that of slow‐growing fish for amino acid metabolism and aerobic capacity [22–32% increase for citrate synthase (CS), aspartate aminotransferase (AAT) and GDH]. The generally higher glycolytic capacities (PK and LDH) in white muscle of fast‐growing fish indicated higher burst swimming capacities and hence better access to food.     

Spring climate and summer otolith growth in juvenile Arctic charr, Salvelinus alpinus

The influence of different climate variables on the first four years of otolith growth in Salangen Arctic charr, Salvelinus alpinus, was studied over the period 1939?C2005. Salangen is a coastal, low altitude, subarctic lake system located in northern Norway. Climate data, including water temperature, air temperature, ice-cover and precipitation, were available for most of the 67?year period. Water temperatures in May and June had a significant effect on otolith growth during the second growth year, while no relationship between otolith growth and climate variables was found for the first, third and fourth years of otolith growth. Otolith increment size during the third and fourth growth year was autocorrelated with growth during the previous year. Spring snow fall and timing of ice break-up had an indirect effect on growth, as these variables were highly correlated with spring water temperatures. High variation in otolith growth within years and among individuals suggests that individual and age-specific variations in spatial habitat use may confound the direct effects of changing air temperatures and time of ice break-up.     

Diel and seasonal locomotor activity patterns in Arctic charr, Salvelinm alpinus (L.)

One-year-old Arctic charr, Sulvelinus alpinus (L.), of the Hornavan strain were tested from February 1985 to January 1986 in an attempt to get an increased understanding of the annual rheotactic behaviour as well as the die1 and seasonal locomotor activity pattern. An annular stream tank equipped with photocells was used to measure the direction of swimming movements as well as the number of passings. From February to late May the locomotor activity was low but increased in July and peaked in September. After November the locomotor activity was again at low winter levels. During the activity peak from July to November the majority ofall movements was directed against the current while no preference for direction was noted during the rest of the year. The high level of swimming movements directed against the current in late summer and autumn may be related to an innate habitat change. From February until June, the charr exhibited a bimodal diurnal activity pattern. In July activity was evenly spread over the whole 24- hour period and in August and September activity was again mainly diurnal with a bimodal pattern. In October and November the activity was mainly nocturnal and in December and January activity was concentrated in the short light period. Both annual and die1 activity are discussed in relation to earlier findings in general locomotor activity in Arctic charr and other salmonids.     

Effects of thermal regime on energy and nitrogen budgets of an early juvenile Arctic charr, Salvelinus alpinus, from Lake Inari

The feed intake, growth, oxygen consumption and ammonia excretion of juvenile Arctic charr were measured over period of four weeks at different temperatures which were either constant (11.0, 14.4, 17.7 °C) or fluctuated daily (14.3 ± 1 °C). Maximum feed intake was estimated to occur at 14.3 °C, while oxygen consumption and nitrogen excretion were highest at the highest temperature, and growth rate was estimated to be highest at 13.9 °C. Feed conversion efficiency was estimated to be highest at 13.2 °C, where over 62.7% of ingested energy was allocated to growth. Metabolic rate accounted for 16–30% of ingested energy and nitrogen excretion was under 3% of ingested energy. The nitrogen budget was under similar thermal influences to the energy budget. Thermal fluctuation increased metabolic rate, but not feed intake, leading to a reduction in feed conversion efficiency under fluctuating temperature conditions.     

Effects of food quality on maturation rate in Arctic charr, Salvelinus alpinus (L.)

The effects of food quality on maturation rate were followed in progeny from wild Arctic charr, caught in Lake Fattjaure, northern Sweden. The fish were reared at five different food quality levels. In the first summer, the fry were given feed of three qualities: diluted with cellulose in three proportions (0, 15 and 25%). During the second winter the impact of changed food quality level was studied by transferring half the fish from the high (control) to the low food quality level and vice versa.
Maturation rate was lower in males reared at the high and improved food quality levels than in males reared at the medium, low and reduced food quality levels. The maturation rate in females was similar at all levels, though the rate tended to increase at the reduced food quality level.
Fish reared at the high and medium levels had similar growth rates, whereas fish at the low food quality level experienced slower growth. Reduced food quality did not arrest the growth offish, whereas improved food quality enhanced their growth.     

Diet, gastric evacuation rates and food consumption in a stunted population of Arctic charr, Salvelinus alpinus L., in Takvatn, northern Norway

Diet and food consumption of a stunted population of Arctic charr in Takvatn, northern Norway, was studied throughout the ice-free season. Chironomid pupae dominated the diet in June, July and August, while zooplankton and three-spined sticklebacks dominated in September and October. Estimates of gastric evacuation rates were obtained from laboratory experiments with wild-caught, acclimated charr, and these estimates were used for in situ estimation of food consumption from weights of stomach contents. The daily food consumption was greatest in July, after which it decreased markedly towards autumn. The estimated food intake rates were low, especially in September and October. The food supply appears to be restricted, which is probably the main reason why the charr in Takvaln exhibit stunted growth.     

Exercise,agonistic behaviour and food acquisition in Arctic charr,Salvelinus alpinus

Synopsis Although swimming is energetically costly, a number of studies on salmonid species have demonstrated increased growth rates in fishes forced to swim for prolonged periods at moderate speeds (typically 1–2 body lengths per sec). This suggests that additional energetic costs of swimming are more than met by alternative compensatory gains. The mechanisms underlying such effects are not fully understood. In this paper, we describe an experiment designed to examine one possible mechanism, namely a swimming-induced inhibition of aggression, with consequent beneficial effects on growth. The study used Arctic charr,Salvelinus alpinus, a species for which a positive relationship between exercise and growth has been clearly established. Using direct behavioural observations on small groups, we demonstrate that individuals displaying high levels of aggressive behaviour are able to monopolise access to food and that enforced swimming at a moderate speed (1 body length per sec) reduces the incidence of aggression although not the degree of monopolisation of food shown by aggressive individuals. These results suggest that the enhanced growth rates accompanying enforced swimming may reflect lower energetic costs of reduced aggressive activity rather than improved access to food by subordinates.     

An electrophoretic and biometric study of Arctic charr, Salvelinm alpinus (L.), from ten British lakes

Samples of Arctic charr, Salvelinus alpinus (L.), were collected from the eight Cumbrian lakes containing all the known English populations. Comparative material was obtained from North Wales and southern Scotland. Comparisons were performed using otolith ages, meristic and morphological characters and by assessing allele frequencies for serum esterase and for skeletal muscle malate dehydrogenase. The results confirm that both basins of Windermere contain spring and autumn spawning races of charr. As well as differing genetically and exhibiting differing growth rates, the two races have different mean gill raker numbers and mean gill raker lengths. The latter two variables were used to derive a discriminant function which enables individual charr to be ascribed to the correct race with 94-96% accuracy. Within both principal races there were further significant differences between charr from the north and south basins of the lake and possibly also between different spawning grounds within each basin. Variation between six of the remaining seven Cumbrian populations is significant but generally no greater than that between the Windermere spring and autumn spawners. The exception is the Ennerdale charr which stands out on morphological, meristic and genetic characters, has a rapid growth rate despite its oligotrophic environment, and is a river rather than a lake spawner. The charr from North Wales and southern Scotland were clearly distinct, both from each other and from the English populations.     

Observations on growth rates of Arctic charr, Salvelinus alpinus (L.), reared at low temperature

Food intake and growth of Arctic charr, Salvelinus alpinus , in fresh water was studied at three temperatures 2·9, 8·4 and 13·1° C). Best growth and highest food intake occurred at 13·1°C. The approximate chemical composition was dependent upon rearing temperature, fish reared at the highest temperature depositing large quantities of fat. Fish were later grown on in either fresh or salt water where two growth patterns were observed. In the light of published data for Salvelinus spp., it is suggested that the poorest growth was shown by fish which were incapable of complete adaptation to saltwater conditions.     

Effect of social interactions on growth rates and conversion efficiency of Arctic charr, Salvelinus alpinus L.

Mixing of fish from size-graded groups did not have detrimental effects on subsequent growth performance as compared with control groups. Coefficients of variance for size, within each population, increased with time demonstrating that social interactions were influencing the growth rates of some individuals. Rates of growth of large fish were higher than those of small indicating the growth of the small fish was suppressed below its potential maximum. The results are discussed in relation to those of previous studies and it is concluded that inferences drawn from changes in coefficients of variance have frequently been incorrect.     

Influence of body weight and temperature on growth rates of Arctic charr, Salvelinus alpinus (L.)

When reared for a period of 6 months at a temperature of 10°C Arctic charr, Salvelinus alpinus , increased in weight from 18 g to approximately 135 g. Specific growth rates decreased as the fish increased in size and the relationship between size and growth rate could be described by the equation:
where G _w is specific growth rate and W is fish weight in grams. Temperature effects upon growth were examined using previously published data. Below the optimum growth temperature, the growth rate of a fish of given size could be predicted using the equation:
where T is the rearing temperature.
Rates of growth of Arctic charr were as high as those reported for other salmonid species reared under similar conditions. Preliminary results suggested that growth rates of charr may be lower in salt water than in fresh water.     

Comparison of growth, diet and food consumption of sea-run and lake-dwelling Arctic charr

Sea-run post-smolt Arctic charr Salvelinus alpinus , (15–26 cm) from Storvatn, northern Norway (70°39'48"N) had significantly higher average specific growth rates in two years (1·64 and 1·66) than the corresponding lake-dwelling charr (0·53 and 1·20). The post-smolts displayed fast compensatory growth in the first 2–3 weeks of their sea residency, but then almost stopped growing prior to their return to fresh water. Lake-dwelling charr grew more evenly during the same time period. Thus, the anadromous charr may return to the lake after only 5–6 weeks in the sea, because the potential to maintain a high growth rate in the sea is reduced. The marine diet consisted mainly of the two crustacean plankton species Calanus finmarchicus , and Thysanoëssa , sp. (88%), and less of fish (6%), insects (4%) and benthos (2%). The diet of lake-dwelling charr consisted mainly of insects (58%, mostly chironomid pupae) and zoobenthos (29%), and less of zooplankton (13%) during the same time period. Although post-smolts had the highest growth rates, they had significantly lower food consumption rates and higher frequencies of empty stomachs than the corresponding lake-dwelling fish. Possible explanations for this paradox are discussed in relation to stomach evacuation rates, water temperature, feeding behaviour and the energy content of the food in the two environments.     

Intraspecific competition and density dependence of food consumption and growth in Arctic charr

1. Intraspecific competition for restricted food resources is considered to play a fundamental part in density dependence of somatic growth and other population characteristics, but studies simultaneously addressing the interrelationships between population density, food acquisition and somatic growth have been missing. 2. We explored the food consumption and individual growth rates of Arctic charr Salvelinus alpinus in a long-term survey following a large-scale density manipulation experiment in a subarctic lake. 3. Prior to the initiation of the experiment, the population density was high and the somatic growth rates low, revealing a severely overcrowded and stunted fish population. 4. During the 6-year period of stock depletion the population density of Arctic charr was reduced with about 75%, resulting in an almost twofold increase in food consumption rates and enhanced individual growth rates of the fish. 5. Over the decade following the density manipulation experiment, the population density gradually rose to intermediate levels, accompanied by corresponding reductions in food consumption and somatic growth rates. 6. The study revealed negative relationships with population density for both food consumption and individual growth rates, reflecting a strong positive correlation between quantitative food intake and somatic growth rates. 7. Both the growth and consumption rate relationships with population density were well described by negative power curves, suggesting that large density perturbations are necessary to induce improved feeding conditions and growth rates in stunted fish populations. 8. The findings demonstrate that quantitative food consumption represents the connective link between population density and individual growth rates, apparently being highly influenced by intraspecific competition for limited resources.     

Epigenetic regulation of trophic morphology through feeding behaviour in Arctic charr, Salvelinus alpinus

Several models of speciation suggest that in species that are phenotypically plastic, selection can act on phenotypic variation that is environmentally induced in the earliest stages of divergence. One trait that could be subject to this process is foraging behaviour, where discrete foraging strategies are common. One species which is highly plastic in the expression of phenotype, the Arctic charr, Salvelinus alpinus (L.), is characterized by discrete variation in the anatomy of the head and mouthparts. These traits have been shown to have a functional significance, but the expression of which is thought to be at least partly phenotypically plastic. Here we test the hypothesis that foraging behaviour may regulate the anatomy of the head and mouthparts in Arctic charr. In a dyad experiment, size‐matched pairs of fish from a mixed family group were fed a diet of either Mysis (a hard‐bodied shrimp) or Chironomid larvae. Nine morphometric measures of head dimensions that describe wild trophic morphs were measured at the start of the experiment and 24 weeks later. Principal component scores of size‐corrected morphometric measures showed highly significant differences between fish exposed to the two diets. Univariate ANOVA analysis of the head morphometric variables showed that fish fed on Chironomids developed longer, wider jaws, longer heads and a larger eye for a given body length than did those fish fed upon Mysis. We conclude that foraging anatomy in Arctic charr is phenotypically plastic and that variation in foraging behaviour that results in feeding specialization in the wild could induce variation in head anatomy. This in turn could reinforce foraging specialization. Very rapid epigenetic divergence into distinct feeding morphs (as demonstrated here) would allow selection to act at more than one mode and thus could promote rapid evolutionary divergence, initially prior to genetic segregation, in species which are highly plastic. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 43–49.     

Effects of the eye fluke Diplostomum spathaceum on the energetics and feeding of Arctic charr Salvelinus alpinus

In the present study, effects of the trematode eye fluke Diplostomum spathaceum infection and parasite‐induced cataract on energetic costs and feeding of hatchery‐bred one‐summer‐old Lake Saimaa Arctic charr Salvelinus alpinus were examined. The energetic effects (measured as oxygen uptake) of acute eye fluke infection on S. alpinus were negligible, but in the chronic infections mean oxygen consumption of infected fish was significantly higher than that of controls. Feeding capability of S. alpinus was studied in aquarium experiments, and uninfected healthy‐eyed fish reacted to zooplankton more rapidly and caught zooplankton with better success compared to individuals with cataracts in the same school. The results suggested that to compensate energetic costs and impaired feeding capability caused by eye fluke, cataract‐bearing fish spend more time foraging, which may result in increased predation risk.     

An investigation of factors controlling food intake in Arctic charr, Salvelinus alpinus L.

Digestibility values were determined for diets of varying protein content fed to Arctic charr. The use of these values, in conjunction with food intake data made possible the study of the effects of various parameters on the control of food consumption. Dietary energy content may be more important than protein content in regulating consumption. The regulation of food intake may be partially mediated through factors influencing gastric emptying coupled with mechanisms monitoring dietary metabolizable energy.     

Effects of crude rapeseed oil on lipid composition in Arctic charr Salvelinus alpinus

This study investigated the effects of crude rapeseed oil (RO) on lipid content and composition in muscle and liver of Arctic charr Salvelinus alpinus . Triplicate groups were fed diets containing fish oil (FO):RO ratio of 100:0, 75:25, 50:50 and 25:75 until two-fold mass increase. Total lipid content increased significantly in the liver with higher proportion of RO in the diet. Profound effects were seen in the fatty acid composition in the analysed tissues with a reduction in 20:5n-3 and 22:6n-3 and an increase in 18:2n-6 with higher RO content in the diets. A drop in cholesterol content was seen at 25% inclusion of RO in both tissues. Wild-caught fish contained a considerably higher amount of 20:4n-6 in both storage and membrane lipids of white muscle compared with the experimental fish.