Temperature preference of juvenile Arctic charr originating from different thermal environments

Temperature preference of juvenile (age 1+) Arctic charr (Salvelinus alpinus L.) originating from four arctic and sub-arctic populations (Svalbard and mainland northern Norway), representing a range of habitats with different temperature conditions, was studied by use of a shuttle-box system which allowed individual fish to control their environmental temperature. Based on the assumption that adaptations to long-lasting differences in thermal environments would affect temperature preference, we expected that Arctic charr from the high arctic Svalbard would prefer a lower temperature than the charr from two well-studied sub-arctic mainland lakes (i.e. one anadromous charr population from Storvatn, Hammerfest and two sympatric resident charr morphs from Fjellfrøsvatn, Målselv). There were, however, no significant differences in temperature preference among the four populations after 24 h exposure to the shuttle-box system, although the charr from the omnivore upper-water sympatric morph of Fjellfrøsvatn used significantly longer time to reach a stable thermal preferendum than the fish of the other populations. The average temperature preference at the end of the trials ranged between 10.9 and 11.6 °C among the populations. The lack of population differences suggests that temperature preference is not a polymorphic trait under strong selection in Arctic charr.     

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.     

Growth and mortality of Arctic charr and European whitefish reared at low temperatures

This comparative study explores how low temperatures affect the mortality and growth of first generation hatchery-reared progeny of subarctic populations of Arctic charr (Salvelinus alpinus L.) and European whitefish (Coregonus lavaretus L.). Replicate fish groups where held under simulated natural light regimes (70°N) at three constant temperatures (1, 3 and 6°C). The mortality of Arctic charr was low (≤1.4%) at all temperature treatments, whereas the mortality of whitefish increased with decreasing temperature from 6% at 6°C to 33% at 1°C. The Arctic charr exhibited higher growth rates than whitefish at all three temperature regimes. All groups of Arctic charr increased in weight, whereas whitefish held at 1°C did not gain weight throughout the experimental period of 133 days. Arctic charr exhibited a large intraspecific variability in growth leading to large variations in size-structure, whereas whitefish in contrast showed very homogenous growth and size-structure patterns; a dissimilarity probably related to species-specific differences in antagonistic behaviour. Evidently, Arctic charr are more cold water adapted than whitefish and are able to maintain growth at extremely low temperatures. Arctic charr thus appear to be the most suitable species for aquaculture at low water temperatures.     

Ecological polymorphism in Arctic charr

Arctic charr, Salvelinus alpinus (L.), commonly exhibits two coexisting morphotypes, dwarf and normal charr, which are characterized by differences in adult body size and colouration. We tested whether or not the morphotypic differences were genetically determined in rearing experiments with offspring of the two morphs and of their crosses. The experiments suggest that this ecological polymorphism in Arctic charr is largely environmentally determined. When reared under similar conditions, offspring of each of the two morphs differed little in size at the same age, and they had the same early developmental rate and maturation pattern. Moreover, the presence of parr marks along the flanks of the fish, one characteristic of dwarf charr, depended on body size and not on parental morph. Genetic differences between the morphs were suggested for growth rate during the second year of life, and for jaw morphology. Comparisons between charr life histories in captivity and in the wild suggest that ecological polymorphism in Arctic charr is chiefly a result of variation in growth conditions between different habitats.     

Parental effects on embryonic viability and growth in Arctic charr Salvelinus alpinus at two incubation temperatures

The parental influences on three progeny traits (survival to eyed‐embryo stage, post‐hatching body length and yolk‐sac volume) of Arctic charr Salvelinus alpinus were studied under two thermal conditions (2 and 7° C) using a factorial mating design. The higher temperature resulted in elevated mortality rates and less advanced development at hatching. Survival was mostly attributable to maternal effects at both temperatures, but the variation among families was dependent on egg size only at the low temperature. No additive genetic variation (or pure sire effect) could be observed, whereas the non‐additive genetic effects (parental combination) contributed to offspring viability at 2° C. In contrast, any observable genetic variance in survival was lost at 7° C, most likely due to the increased environmental variance. Irrespective of temperature, dam and sire–dam interaction contributed significantly to the phenotypic variation in both larval length and yolk size. A significant proportion of the variation in larval length was also due to the sire effect at 2° C. Maternal effects were mediated partly through egg size, but as a whole, they decreased in importance at the high temperature, enabling a concomitant increase in non‐additive genetic effects. For larval length, however, the additive component, like maternal effects, decreased at 7° C. The present results suggest that an exposure to thermal stress during incubation can modify the genetic architecture of early developmental traits in S. alpinus and presumably constrain their short‐term adaptive potential and evolvability by increasing the amount of environmentally induced variation.     

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.     

A comparison of growth patterns between a stunted and two large predatory Arctic charr populations under identical hatchery conditions

Arctic charr are characterized by an extensive variability in growth and body size in natural waters. Although growth traits may involve a significant heritable component, most of this intraspecific variation presumably is environmentally induced and thus attributable to phenotypic plasticity. In the present study, size-at-age and length–weight relationship (body condition) were assessed for three Finnish Arctic charr populations of different geographical origins and extreme size forms (a stunted vs. two large-growing, predatory charr) held under standardized rearing conditions for 3 years (up to 37 months after hatching). In particular, our interest was to investigate whether the differences in growth between the large and the stunted charr as observed in the wild populations would diminish when the fish are offered suitable food in abundance. Population-specific mean body size and condition differed significantly in 0+, 1+, 2+ and 3+ fish. However, the identical rearing conditions resulted in the originally stunted charr reaching a comparable final mean size (317 mm/427 g) as the large charr populations (343 mm/510 g and 359 mm/497 g). Some individuals were of the same size as their parents at spawning already at the age of 0+ years. Furthermore, length–weight regression residuals of the stunted charr developed to a notably high level, indicating the largest final condition mean. The increase of size variation (CV of weight) in stunted charr lasted for over two growth seasons, whereas in large charr it remained stable since the end of the first summer. Variations in mortality and sexual maturation at age 2 seemed to be less relevant factors affecting overall growth performance. The study demonstrates an example of the high plasticity involved in the growth of fish: the stunted charr possess a tremendous capacity for growth in a benign environment, virtually corresponding to that observed in the large predatory populations.     

Latitudinal variation in growth among Arctic charr in eastern North America: evidence for countergradient variation?

Biological data from 66 populations of Arctic charr, Salvelinus alpinus, from eastern North America were analysed to test the applicability of the countergradient hypothesis as an explanation of differences in seasonally adjusted growth rates. Samples were obtained along a 37° latitudinal gradient and partitioned among anadromous, normal lacustrine, and dwarf lacustrine Arctic charr morphotypes. Models relating length-at-age or age-specific growth rates to latitude were estimated for each morphotype. Length-at-age declined with latitude for anadromous and lacustrine charr. Age-specific growth rates also varied with latitude, particularly for normal lacustrine charr. Results of analyses provide support for the countergradient hypothesis in growth performance of normal lacustrine morphotypes, where northern populations compensate for the shorter growth season with a greater rate of growth than southern populations. Anadromous charr exhibited equivocal evidence of countergradient variation, while results for dwarf lacustrine Arctic charr populations were inconclusive owing to the limited range of ages, and latitudes for which data were available.     

The impact of intra- and interspecific interactions on young-of-the-year brook charr, in temperate lakes

The abundance, growth, spatial distribution, and feeding habits of five allopatric brook charr, Salvelinus fontinalis, populations (young-of-the-year, 0+ juveniles; YOY) were compared with five other populations living sympatrically with white sucker, Catostomus commersoni. The study was made in oligotrophic lakes of the Laurentian Shield (Québec, Canada) during three sampling periods in 1989 (July, August and September). The abundance of YOY charr was significantly higher in allopatric than in sympatric populations (45·3 ± 3·8 vs 3·4 ± 3·8 fish/lake caught in 1773 m² of gillnets; P<0·005). The mean length of YOY charr did not differ among allopatric and sympatric populations at each sampling period; July: 60·2 ± 3·0 vs 60·0 ± 4·5 mm; August: 61·9 ± 4·5 vs 63·2 ± 4·1 mm; September: 77·9 ± 8·7 vs 77·3 ± 7·8 mm respectively. Horizontal distribution of allopatric YOY charr did not differ from that of sympatric charr, 65% of the fish being captured within the first 2 m depth and the rest between 2 and 7 m depth. In contrast, the vertical distribution of allopatric YOY charr from both communities was significantly different; 81% of allopatric charr were captured within 0·5 m from the substrate compared to 64% for sympatric charr (P<0·001). Differences in vertical distribution of the fish were related to differences in diet; allopatric charr fed mainly on benthic and large planktonic organisms whereas sympatric charr fed less on these organisms and more on terrestrial organisms. In the lake where YOY charr were most abundant, individuals were spatially segregated into two groups; one ‘littoral’, found in 0–2m depth, and one ‘profundal’, found in 3–6 m depth. Growth, condition, and feeding habits of charr from the two groups were different, especially during the last sampling period.     

Relationships between lake ecology and morphological characters in Icelandic Arctic charr,Salvelinus alpinus

The common occurrence of parallel phenotypic patterns suggests that a strong relationship exists between ecological dynamics and micro‐evolution. Comparative studies from a large number of populations under varying sets of ecological drivers could contribute to a better understanding of this relationship. We used data on morphology of arctic charr (Salvelinus alpinus) and ecological factors from 35 Icelandic lakes to test the hypothesis that morphological patterns among monomorphic charr populations from different lakes are related to interlake variation in ecological characteristics. There is extensive phenotypic diversity among populations of Icelandic charr, and populations are easily distinguished based on overall body morphology. The results obtained in the present study showed that the morphological diversity of charr was related to large‐scale diversity in lake ecology. Variation in charr morphology was related to water origin (e.g. spring fed versus run‐off), bedrock age, and fish community structure. The present study shows how various ecological factors can shape the biological diversity that we observe. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 761–771.     

A morphometric method for estimation of total lipid level in live Arctic charr: a case study of its application on wild fish

Linear body measurements were made on wild Arctic charr Salvelinus alpinus parr (100–200 mm L_F) from two populations in northern Norway during spring (May), summer (June and July) and autumn (October). To reduce handling stress, a method was developed were dorsal and ventral body measurements could subsequently be taken easily from a single picture of the anaesthetized fish. Multiple regression analyses investigated the possible correlation between the body measurements and both total and percentage lipid content of the fish. For both populations and during all seasons, the regression equations gave better estimates for total lipid than for percentage lipid. The regression equations for total lipid accounted for between 67·7 and 89·5% of the variance in lipid content for the different seasons within the lakes. The pooled data within each lake accounted for 62·9 and 81·0% of the variance in total lipid, while the pooled data for both lakes accounted for 67·7% of the variance. In general, the condition factor alone did not give a satisfactory estimate of lipid content of the Arctic charr (r² = 0·003–0·521). Shrinkage on validation values was high (0·20–0·52). Employing a ridge regression method resulted in models with lower r² values and lower shrinkage values (indicating more stable models). Published equations used for hatchery‐reared Arctic charr could not be used on wild fish from the same population. The close correlations between actual and predicted lipid levels found in this study show that morphometric measurements can predict total lipid levels in wild Arctic charr with reasonable accuracy. The most accurate results were obtained when equations were derived from each data set. Therefore, the method has the potential within a single study to estimate lipid levels in live fish as long as some fish can be sacrificed to develop a unique regression equation for each population or experiment.     

Thermal growth performance of juvenile brown trout Salmo trutta: no support for thermal adaptation hypotheses

Using thermal growth data from eight populations of anadromous and lake-feeding brown trout Salmo trutta , hypotheses of adaptation to local optima and countergradient variation in growth were tested. The adaptation to local optima hypothesis suggests that natural selection can shift optimal performance temperatures to match the prevailing temperature in a new or changed thermal niche. In contradiction, the countergradient variation hypothesis suggests that populations from hostile environments perform better than conspecifics from benign environments at all temperatures. In this study, growth capacity varied between populations but there was no significant correlation between any of the estimated thermal performance parameters ( e.g. lower and upper thermal growth limits, optimal temperature for growth and maximum growth capacity) and natural climatic conditions among populations. Hence, S. trutta growth response to temperature lends no support for either of the two suggested thermal adaptation hypotheses. Instead, growth capacity among populations tended to correlate positively with female size at maturity.     

A comparative study of the relationship between light intensity and feeding ability in brown trout (Salmo trutta) and Arctic charr (Salvelinus alpinus)

1. Field observations indicate that the ability to feed at different light intensities may differ between brown trout and Arctic charr, and this is the first study to test this experimentally. To establish a background level of feeding in daylight at midday, trout and charr in two size groups were kept in tanks (one fish per tank) at three constant temperatures (5.0, 10.8 and 13.0 °C) and each fish was offered, one at a time, 50 freshly killed shrimps (Gammarus pulex), the number eaten being recorded. Shrimps could only be taken in the water column because a metal mesh prevented access to dead shrimps on the tank bottom. In a first series of experiments, individual fish were kept at one of 10 natural light intensities (range 0.001–50 lx). In a second series, conditions were similar except that the fish tank was covered in black polyethylene and had a light‐tight lid with a shutter so that light levels could be kept constant, using artificial illumination. In a third series, the fish were fed in total darkness, but the false bottom was removed, allowing access to dead shrimps on the tank bottom as well as in the water column. 2. The results of the first and second series differed interspecifically but were very similar intraspecifically, with no significant differences between the food intake for the two size groups or in the experiments at 10.8 and 13.0 °C. Food intake remained fairly constant at light intensities between 50 lx (dusk or dawn) and 0.03 lx and was similar to that of fish feeding at midday. At 10.8 and 13.0 °C, food intake between 0.03 and 50 lx was higher for trout than for charr, mean values for shrimps eaten per fish being 39.9 for trout (range 36–44, n = 100 fish) and 32.0 for charr (range 28–38, n = 100), but at 5.0 °C, the situation was reversed with mean values of 15.1 for trout (range 11–18, n = 50 fish) and 19.8 for charr (range 17–22, n = 50). 3. As light intensity decreased from 0.04 to 0.001 lx, feeding rate decreased exponentially but was always higher for charr than for trout, with a mean number of shrimps eaten at 0.001 lx of 9.3 for trout (range 5–13, n = 20 fish) and 13.6 for charr (range 9–20, n = 20) at 10.8 and 13.0 °C, and 2.0 for trout (range 1–4, n = 10 fish) and 5.5 for charr (range 2–8, n = 10) at 5.0 °C. In total darkness (false bottom fitted), none of the 50 shrimps was taken by either species. When the false bottom was removed in the third series, the mean number of shrimps consumed over 24 h was eight for trout (range 3–11, n = 20 fish) and 14.9 for charr (range 9–20, n = 20) at 10.8 and 13.0 °C, and two for trout (range 0–4, n = 10 fish) and five for charr (range 3–8, n = 10) at 5.0 °C. 4. Therefore, the feeding ability of trout was superior to that of charr when using photopic vision in daylight and mesotopic vision at dusk and dawn, but inferior to that of charr when using scotopic vision at low light intensity. Charr were also superior at low temperatures and when foraging for food in total darkness. Therefore, as light intensity decreases after dusk in their natural habitat, the advantage in feeding will shift from trout to charr, with the reverse occurring as light intensity increases after dawn.     

The size structure of lacustrine Arctic charr (Pisces: Salmonidae) populations

The size structure of Arctic charr (Salvelinus alpinus) populations was analysed using published information; 44% of populations were bimodal, with both large (normal) and small (dwarf) morphs occurring within a cohort. The remaining populations were unimodal, consisting of normal-sized or of stunted adults. Bimodal populations increased in frequency with latitude and were characteristic of large, deep lakes with few fish species. The age and size at which bimodality developed, the size difference between the morphs, and the frequency of cannibalism in charr populations increased with latitude. A variety of genotypic and phenotypic explanations are examined. The evidence for specific differences between the morphs is unconvincing. Various competition and predation hypotheses fail to explain the occurrence of bimodality. Cannibalism does not cause bimodality despite being strongly associated with it since bimodality develops before charr become cannibalistic. Much of the variation in charr size structure is suggested to be a consequence of increased seasonality in food supply in more northerly environments, coupled with feeding size thresholds. The latter result in larger members of a cohort being able to maintain growth rates on seasonally abundant prey while smaller individuals which cannot catch these items form a second mode of more slowly growing fish. Bimodality is documented in a number of other, predominantly northern, fish species.     

Inter- and intra-population variations in diapause attribute of the two-spotted spider mite,Tetranychus urticae Koch,in Japan

Populations of the two-spotted spider mite, Tetranychus urticae Koch collected from various localities and from various host plants in Japan showed wide variations in diapause attribute. Diapause percentages at 18°C/9L15D varied from nearly 100% in the north to 0% in the south-west. At intermediate latitudes the mites showed wide inter-population variations. Populations on herbaceous hosts in vinyl- or glass-houses gave significantly lower incidence of diapause than those on roses and deciduous fruit trees. Presence of winter hosts and better host quality under protected environments seemed to favour non-diapausing mites. The temperature threshold for diapause expression also varied widely among local populations. Northern populations consistently had higher and less variable thresholds than populations at intermediate latitudes with thresholds between 15 and 18°C. Inbred lines derived from a population in Kyoto exhibited a wide variation in diapause percentage at 18°C. These results show that diapause in T. urticae is a quantitative threshold trait and that populations in central Japan consist of a variety of genotypes with different diapause traits. This might provide a genetic source for adaptation to local and temporal variations in environmental conditions.     

Upper thermal tolerance of wild‐type,domesticated and growth hormone‐transgenic coho salmon Oncorhynchus kisutch

In coho salmon Oncorhynchus kisutch, no significant differences in critical thermal maximum (c. 26·9° C, C_Tmax) were observed among size‐matched wild‐type, domesticated, growth hormone (GH)‐transgenic fish fed to satiation, and GH‐transgenic fish on a ration‐restricted diet. Instead, GH‐transgenic fish fed to satiation had significantly higher maximum heart rate and Arrhenius breakpoint temperature (mean ± s.e. = 17·3 ± 0·1° C, T_AB). These results provide insight into effects of modified growth rate on temperature tolerance in salmonids, and can be used to assess the potential ecological consequences of GH‐transgenic fishes should they enter natural environments with temperatures near their thermal tolerance limits.     

Winter ecology of Arctic charr (<Emphasis Type="Italic">Salvelinus alpinus</Emphasis>) and brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) in a subarctic lake,Norway

We studied habitat choice, diet, food consumption and somatic growth of Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) during the ice-covered winter period of a subarctic lake in northern Norway. Both Arctic charr and brown trout predominantly used the littoral zone during winter time. Despite very cold winter conditions (water temperature <1°C) and poor light conditions, both fish species fed continuously during the ice-covered period, although at a much lower rate than during the summer season. No somatic growth could be detected during the ice-covered winter period and the condition factor of both species significantly declined, suggesting that the winter feeding rates were similar to or below the maintenance requirements. Also, the species richness and diversity of ingested prey largely decreased from summer to winter for both fish species. The winter diet of Arctic charr <20 cm was dominated by benthic insect larvae, chironomids in particular, and Gammarus lacustris, but zooplankton was also important in December. G. lacustris was the dominant prey of charr >20 cm. The winter diet of brown trout <20 cm was dominated by insect larvae, whereas large-sized trout mainly was piscivorous, feeding on juvenile Arctic charr. Piscivorous feeding behaviour of trout was in contrast rarely seen during the summer months when their encounter with potential fish prey was rare as the small-sized charr mainly inhabited the profundal. The study demonstrated large differences in the ecology and interactions of Arctic charr and brown trout between the winter and summer seasons.     

Stable isotope-based determinations of the average temperatures experienced by young-of-the-year Svalbard Arctic charr (Salvelinus alpinus (L.))

Mean ambient water temperatures experienced by individual young-of-the-year (YOY) Arctic charr, Salvelinus alpinus (L.), from a Svalbard lake were estimated using measurements of oxygen stable isotopes (δ¹⁸O) derived from fish otoliths. Otolith-derived water temperatures differed significantly from temperatures recorded at the outlet river of the Dieset watercourse but were consistent with temperatures previously recorded in shallow littoral areas of other Svalbard lakes where YOY charr are commonly found. This indicates that fixed-point monitoring does not necessarily represent the temperatures and thermal habitats used by individual fish. Otolith-derived water temperatures were also positively related to fish length-at-capture and otolith size, although much of the variation remained unexplained. Differences among individuals could be related to variability in food availability and food intake as well as variation in the initial fish size at hatching. Implications for subsequent investigations into how YOY charr respond and adapt to future climate change are discussed.     

Effects of population,family, and diet on craniofacial morphology of Icelandic Arctic charr (Salvelinus alpinus)

We evaluated hypotheses of intralacustrine diversification and plastic responses to two diet environments in Icelandic Arctic charr (Salvelinus alpinus). Full‐sib families of progeny of wild polymorphic charr from two lakes where morphs vary in their degree of phenotypic and ecological divergence were split, with half of the offspring reared on a benthic and half on a limnetic type of diet to estimate family norms of reaction. We focused on variation in craniofacial traits because they are probably functionally related to diet and complement a previous study of body shape in these charr. A hierarchical analysis of phenotypic variation between lakes, pairs of morphs within each lake, and two families within each morph found that phenotypic variation partitioned between families relative to morphs was reduced in the more ecologically diversified population, which is consistent with adaptive diversification. The effect size of plastic responses between lake populations was similar, suggesting little difference in the degree of canalization in contrast to a previous analysis of body form plasticity. Thus, the role that plastic morphological responses play in the adaptive diversification of morphs and different lake populations of Arctic charr may depend on the trait. © 2013 The Linnean Society of London     

Temperature influence on Arctic charr (Salvelinus alpinus L.) antibody response to a cellular antigen

In order to elucidate the immune responses of Arctic charr in relation to temperature, groups were acclimated to a moderate (9°C) and a cold temperature regime (4°C), as well as subjected to a temperature decrease (from 9 to 4°C) immediately prior to an immunization with sheep red blood cells. The charr kept at 9°C responded with increased primary and secondary antibody titres, as seen by direct haemagglutination, while fish at 4°C, as well as the fish subjected to a temperature reduction, displayed lower and lowest antibody titres, respectively, and only after a second immunization. It is concluded that Arctic charr can respond to a cellular antigen with a humoral immune response typical for other teleosts, but that the immune response is delayed and diminished at low temperatures. This temperature-induced immune suppression is intensified if the fish have not been acclimated to cold water prior to immunization. Accepted: 10 October 1999