Seasonal changes in sea-water tolerance of Arctic charr (Salvelinus alpinus)

Summary Groups of Arctic charr,Salvelinus alpinus, which had been acclimated to water with a salinity of 7 g·l^–1 and natural temperature and photoperiod, were exposed to water with different salinities and temperatures in June, September and February. At a salinity of 15 g·l^–1, plasma osmolality, plasma Na⁺, Cl^–, Mg²⁺ concentrations and the activity of gill Na-K-ATPase were stable, irrespective of temperature and season. In June, the charr were able to regulate blood plasma ionic levels within narrow limits when exposed to a salinity of 34 g·l^–1 (sea water) and a temperature of 8°C. The hypo-osmoregulatory capacity was less, but sufficient if the temperature was only 1°C during the seawater exposure. At the start of the experiment, the gill Na-K-ATPase activity was significantly higher in June than corresponding enzyme activities in September and February. Furthermore, an increase in gill Na-K-ATPase activity during the seawater exposure (8°C) was seen in June. Irrespective of ambient temperature and salinity, no fish died during the June experiments. In September and February, exposure to sea water produced marked increases in plasma osmolality and plasma ion concentrations. There were no changes in gill Na-K-ATPase activity. Consequently, the fish became dehydrated and were moribund after a short period of seawater exposure. Highest mortality was recorded when charr were exposed to winter sea conditions (34 g·l^–1 and 1°C) in February. The results indicate that an increase in daylength induce a hypo-osmoregulatory capacity in the Arctic charr during summer. In fall and winter, however, reduced daylength are accompanied by poor hypo-osmoregulatory capacity. This leads to high mortality as a result of increased electrolyte levels and dehydration.     

Seasonal variation in the temperature preference of Arctic charr (Salvelinus alpinus)

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Seasonal variation in thermal preference of Arctic charr from a North Norwegian population (70°N) was studied in an electronic shuttlebox, which allowed the fish to control the water temperature by moving between two chambers. The fish were acclimated to 12 °C and a natural photoperiod before measurements. Fish were tested in autumn (September–October), winter (January–February), spring (April–May) and summer (June–August).     

Seasonal and ontogenetic variations in resource use by two sympatric Arctic charr morphs

The study compares the resource utilization of two sympatric Arctic charr morphs over an annual period in a subarctic lake. The two morphs are reproductively isolated in time and place of spawning, and are referred to as the littoral and profundal morphs (L-morph and P-morph) according to their spawning habitats. Fish were sampled monthly (ice-free season) or bimonthly (winter) using gillnets in the main lake habitats. The spatial range of the P-morph was restricted to the profundal zone throughout the whole annual period. The L-morph in contrast utilized all main habitats, exhibiting distinct seasonal and ontogenetic variations in habitat distribution. In the spring, the whole L-morph population was located along the bottom profile of the lake, in profundal and littoral habitats. During summer and autumn, habitat segregation occurred between different life-stages, juveniles mainly utilizing the profundal, pre-adults the pelagic and adult fishes the littoral zone. During winter the whole population was assembled in the littoral habitat. The L-morph also had large seasonal and ontogenetic variations in their feeding ecology, with littoral zoobenthos, zooplankton and surface insects being important prey. The P-morph had a narrower diet niche mainly consisting of chironomid larvae and other profundal zoobenthos. Hence, the two Arctic charr morphs exhibited a consistent resource differentiation during all annual seasons and throughout their life cycles, except for a dietary overlap between P-morph and juvenile L-morph charr in the profundal during summer. The findings are discussed in relation to resource polymorphism and incipient speciation.     

Food borne parasites as indicators of trophic segregation between Arctic charr and brown trout

The habitat and diet choice and the infection (prevalence and abundance) of trophically transmitted parasites were compared in Arctic charr and brown trout living sympatrically in two lakes in northern Norway. Arctic charr were found in all main lake habitats, whereas the brown trout were almost exclusively found in the littoral zone. In both lakes the parasite fauna reflected the niche segregation between trout and charr. Surface insects were most common in the diet of trout, but transmit few parasites, and accordingly the brown trout had a relatively low diversity and abundance of parasites. Parasites transmitted by benthic prey such as Gammarus and insect larva, were common in both salmonid host species. Copepod transmitted parasites were much more common in Arctic charr, as brown trout did not include zooplankton in their diets. Parasite species that may use small fish as transport hosts, were far more abundant in piscivorous fish, especially brown trout. The seasonal dynamics in parasite infection were also consistent with the developments in the diet throughout the year. The study demonstrates that the structure of parasite communities of charr and the trout is highly dependent on shifts in habitat and diet of their hosts both on an annual base and through the ontogeny, in addition to the observed niche segregation between the two salmonid species.     

A Comparison of Floy and Soft VIalpha tags on Hatchery Arctic Charr, with Emphasis on Tag Retention, Growth and Survival

The Floy FTF-69 'fingerling' tag and the soft VIalpha tag are designed to be used on small salmonid fishes (> 100 and > 150mm, respectively). The two tags were compared for 160 days on hatchery-reared Arctic charr, Salvelinus alpinus, (170–209mm). Tag retention and effects on growth rate and survival were analysed. VIalpha-tagged and untagged control fish had similar growth rates and grew significantly faster than Floy-tagged fish during the experiment. Tag retention was significantly higher for Floy tags (94%) than for VIalpha tags (78%). Most VIalpha tag loss (11%) took place within the first 10 days of tagging while there were no Floy tags lost during this period. Survival rates were about equal for all three groups (96–98%), and thus the tags did not seem to affect the survival of charr under hatchery conditions.     

The relationship between the digestive enzymes in arctic char,Salvelinus alpinus (Salmonidae,Osteichthyes) and its ability to survive in extreme environments

Digestive enzymes of chars from seven Austrian lakes have been studied. Trypsin activity ml⁻¹ fluid is proportional to body size but inversely proportional to the amount of food in the digestive tract. The pancreas appears to secrete trypsin ± continuously and the activity measureable in the intestinal fluid seems to depend largely on the gut passage rate. There is no seasonal rhythm of enzyme activity. Amylase activity is very weak. Pepsin activity seems to depend mostly on the dilution of the gastric juice. The results indicate that the steering mechanisms of digestive enzymes in salmonidae are well distinct from other families. This work was part of a PH. D. thesis (Reimer, 1984). This work was part of a PH. D. thesis (Reimer, 1984).     

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.     

Seasonal dynamics and persistence of Gyrodactylus salaris in two riverine anadromous Arctic charr populations

The seasonal occurrence of the monogenean ectoparasite Gyrodactylus salaris Malmberg infecting Arctic charr, Salvelinus alpinus (L.) in the two rivers Skibotnelva and Signaldalselva in northern Norway was studied in the period from autumn 2003 to autumn 2005. Skibotnelva has been infected with the parasite since 1979, and treated with rotenone twice. Most likely resident Arctic charr avoided the rotenone treatment in small tributary streams, and thus was the source of the repeated re-infection of this river. G. salaris was first recorded in Signaldalselva in the year 2000 and it is still untreated. Unlike Atlantic salmon, which is highly susceptible to G. salaris, Arctic charr can display a wide range of host-responses to G. salaris infections. Arctic charr were sampled by electro fishing with a total sample of 681 Arctic charr. The results from this study demonstrate an evident seasonal dynamic in G. salaris infection in charr in both rivers. Parasite intensities fluctuated with the rise and fall in temperature through the year, with an autumn high and spring low. There was a significantly lower prevalence and mean intensity of G. salaris in Skibotnelva than in Signaldalselva. There were also a lower prevalence and intensity of G. salaris in the older than in the youngest charr. The different history of infection and treatment in the two rivers might be the underlying cause of these observed dissimilarities. The current study indicates that Arctic charr is a good natural host for G. salaris.     

Characterisation and expression of the paired box protein 7 (Pax7) gene in polymorphic Arctic charr (Salvelinus alpinus)

Arctic charr (Salvelinus alpinus L.) from Lake Thingvallavatn, Iceland occur as four distinct morphs: large benthivorous (LB), dwarf benthivorous (DB), piscivorous (PI) and planktonivorous (PL). The morphs differ with respect to body size, head morphology, growth rate, and life history. The aim of this study was to investigate the paired box protein 7 (Pax7) gene as a candidate for such polymorphisms due to its importance in cranio-facial, skeletal muscle, and central nervous system development. No variation in coding and intronic sequences was found between morphs. We identified 10 alternate Pax7 isoforms with insertions/deletions: a four-residue (GNRT) deletion, a GEASS insertion truncated by the first serine residue (GEAS), and a thirteen-residue insertion (GQYA/TGPEYVYCGT). The latter insertion with a threonine (T) contains a putative casein kinase II (CK-2) phosphorylation site. Pax7 spatial expression patterns were identical in embryos of DB-, LB-, and PL-morphs, and were similar to those described for zebrafish Pax7c, but a difference in temporal expression for segmentation was observed between DB and LB morphs. At the end of segmentation, novel expression was observed in the mandibular region as two bilateral domains. The potential role of multiple alternative splicing of the Pax7 gene for the generation of different Arctic charr morphs is briefly discussed.     

Thermal habitat use and growth in young-of-the-year Arctic charr from proximal fluvial and lacustrine populations in Labrador,Canada

Predicted increases in temperature associated with climate change are expected to have consequences for fish, in particular for Arctic charr, Salvelinus alpinus, a cold-adapted fish species. Despite differences in predicted hydroecological responses to climate change in fluvial and lacustrine environments, little is known of whether fluvial and lacustrine Arctic charr populations may respond differently to increasing temperatures. In order to compare growth and thermal habitat use between habitat types, otolith-inferred average water temperatures estimated from whole otoliths and fork lengths at capture were measured for young-of-the-year (YOY) Arctic charr obtained from two proximal fluvial and lacustrine sites in Labrador, Canada. Otolith-inferred average experienced water temperatures were not significantly correlated with air temperatures at both sites, suggestive of behavioural thermoregulation by YOY. The majority of Kogluktokoluk Brook (fluvial) YOY were found using water temperatures consistent with laboratory determined preferred temperatures for juvenile Arctic charr, whereas most Tom's Pond (lacustrine) YOY were found using temperatures ranging between preferred temperatures and optimal temperatures for growth. There was no consistent difference between mean water temperatures used between YOY from the two sites. Otolith-inferred average experienced water temperatures were only correlated to fork lengths in Tom's Pond YOY. The lack of correlation in Kogluktokoluk Brook YOY is argued to reflect resource partitioning occurring as a result of territoriality known to occur among stream salmonids. The limited range of temperatures used by fluvial YOY in this study, particularly the lack of cooler temperatures, also suggests that fluvial YOY may face barriers to thermal refugia, and as a result may be particularly vulnerable to climate change.     

Reproduction,condition and food of adult arctic charr (Salvelinus alpinus,L.) in Lake Pavin (Massif Central,France)

RésuméTo determine the role and the importance of adult fish predation directly and indirectly in the food web of oligomesotrophic Lake Pavin, reproduction, condition, and feeding activity of the dominant adult chctic charr (Salvelinus alpinus, L.) were studied. Fish were caught by gillnets of different mesh sizes once a month from April to December 1992. The gonad cycle showed that the spawning period occurred between October to December. Relative fecundity is about 2509 ovules per kg of female fish and ovule mean diameter is 4.7 mm ( 0.6 mm). Seasonal variations of the sex-ratio (Max. in April = 2.67 and Min. in October=0.19) indicated the intralake migrations of females and males. The condition coefficients showed higher values for females than males; the maximum was observed in summer for both sexes and the minimum in April and in October for males and females, respectively. Low values of the condition were also observed during the spawning period. High feeding activity was recorded in spring and in December after reproduction; low feeding activity was observed in October during reproduction. Asellus aquaticus was quantitatively the dominant prey from April to September; during the sp awning period, S. alpinus consumed its own eggs, particularly in large quantity in December. Daphnia longispina was numerically the dominant prey all over the studied cycle and the electivity (Ivlev Index) of arctic charr on the latter prey ranged from +s+0.79 to +0.16. During April, more than 15 500 individuals of D. longispina per kg of fish were found in the gut contents. The selective predation of S. alpinus on benthic (A. aquatics) and zooplankton prey (D. longispina) indicated that the fish immigrate during their feeding activity from benthic/littoral to the pelagic area.Afin de d terminer le rôle et l'importance de la prédation directe et indirecte de l'omble chevalier adulte (Salvelinus alpinus, L.) dans le réseau trophique du lac Pavin, milieu oligo- mésotrophe, nous avons étudié la reproduction, deux facteurs de condition et l'activité alimentaire de ce poisson qui est l'espéce dominante dans ce lac. Les poissons ont été capturés par des filets maillants de différent vide de maille selon un pas d'échantillonnage mensuel d'avril á décembre 1992. Le cycle du développement des gonades montre que la période de reproduction se déroule entre octobre et décembre. La fécondité relative calculée est de 2509 ovules par kg de femelles et le diamétre moyen d'un ovule est de 4,7 mm ( 0,6). Les variations saisonniéres du sex-ratio (Max. en avril = 2,67 et Min. en octobre=0,19) indiquent le déplacement actif et successif des femelles et des mâles. Les variations saisonniéres des coefficients de condition indiquent une copulence plus importante des femelles par rapport aux mâles avec des valeurs maximales en été pour les deux sexes et des valeurs minimales en avril et en octobre pour les mâles et les femelles respectivement. De faibles valeurs des facteurs de condition ont été observées pendant la période de reproduction pour les deux sexes. L'activité alimentaire de l'omble chevalier est intense au printemps et au mois de décembre aprés la fraie et faible au mois d'octobre pendant la reproduction. Asellus aquaticus est, en termes de biomasse, la proie dominante d'avril á septembre. Pendant la période de reproduction, S. alpinus se nourrit de ses propres oeufs sur les zones de frayéres et en décembre la quantité d'oeufs consommée par le Salmonidé est importante. Daphnia longispina est, en termes d'abondance, la proie dominante tout au long de cette étude et l'indice d'électivité (Indice d'Ivlev) de l' ombre chevalier vis-á-vis de cette proie varie entre + 0,79 et + 0,16.De plus, au mois d'avril, plus de 15 500 D. longispina par kg de poisson ont été enregistrées dans les contenus stomacaux. La prédation sélective de S. alpinus sur des proies benthiques (A. aquaticus) et zooplanctoniques (D. longispina) indique que le poisson effectue des déplacements en fonction de son activité alimentaire de la zone littorale/benthique á la zone pélagique.     

Life-history styles and somatic allocation in iteroporous arctic charr and semelparous pink salmon

Synopsis RNA/DNA ratios were used to estimate recent somatic allocation of spawning arctic charr,Salvelinus alpinus, and pink salmon,Oncorhynchus gorbuscha. Smaller arctic charr morphs had lower ratios than larger morphs. Larger male pink salmon had lower ratios than smaller males. Thus recent somatic allocation (anabolic metabolic activity and growth) is coupled with cumulative lifetime growth in iteroparous arctic charr, but is uncoupled in semelparous pink salmon. These data suggest that somatic investment in these species is being differentially effected by the energetic costs of reproduction.     

Diel and density-related changes in food consumption and prey selection by brook charr in a New Hampshire stream

Synopsis We report the results of a field study testing influences of both density and changes over the diel cycle on food consumption and prey selection by brook charr,Salvelinus fontinalis. Charr density in replicate 35 m long sections of a New Hampshire stream was adjusted to either medium or high levels (relative to natural densities). Diets of charr and the availability of drifting prey were then sampled every four hours for 24 hours. There were no significant diel changes in the weight of prey consumed by charr per four hours, though there was some indication of reduced feeding at night. Chary fed selectively on different prey taxa, showing most preference for cased caddis larvae. Several species of mayflies and stoneflies were selected more strongly during the day than at night. Charr fed selectively on larger prey during the daytime but showed no size-selection at night. The density of charr had no significant effects on either their rate of food consumption or on selection for prey of different taxa or sizes.     

Adaptations to Stochastic Environmental Variations: The Effects of Seasonal Temperatures on the Migratory Window of Svalbard Arctic Charr

The circumpolar Arctic charr, Salvelinus alpinus, is ideal for studying how environmental factors affect life history in fishes. Charr populations demonstrate a tremendous ecological plasticity and adaptations to harsh environments. Arctic charr is the only freshwater fish on Svalbard, including anadromous, resident, and landlocked stocks. Freshwater lake systems on Svalbard are characterized by very low water temperatures, long-term or even permanent ice cover, and low levels of nutrients. Food is thus limited and may lead to growth stagnation and early maturity in Arctic charr. The individual growth pattern may alternatively follow a sigmoid-shaped curve, caused by a shift to either cannibalistic or anadromous (migration to sea) behaviour. In lake systems that include migratory charr, the population may consist of a mixture of parr, postsmolt, and adult migratory individuals, as well as small-sized resident, large-sized resident (cannibals), and large formerly resident individuals transformed to anadromy. Our study in the Lake Dieset watercourse (79°N), Svalbard, demonstrates that the annual water flow in the outlet river is strongly correlated to air temperatures and provides a passage to the sea, allowing the charr access to the nutrient-rich seawater environment, during at most two months each year. During one of the years studied, the youngest and small-sized part of the sea-going stock was prevented from ascending the river and probably suffered mortality during winter. The migratory window of the Arctic charr in Lake Dieset is therefore highly variable among years and thus unpredictable. We hypothesize that in worst case scenarios (cold years, low water discharge), climatic variations may occasionally prevent charr from migrating upstream in Svalbard lake systems in late autumn, resulting in high mortality in the population.     

Seasonal changes in hypoosmoregulatory ability in landlocked and anadromous populations of Arctic charr,Salvelinus alpinus,and Atlantic salmon,Salmo salar

Synopsis Seasonal changes in hypoosmoregulatory ability were compared in landlocked and anadromous strains of Arctic charr and Atlantic salmon. Seawater adaptability was assessed using periodic 48 h seawater challenge tests with 25. seawater. The landlocked strains of Arctic charr, two from northern Sweden and one from Southern Norway, displayed similar seasonal changes in seawater adaptability as the anadromous strain. Seawater tolerance increased during spring and remained high until the end of July — early August after which it declined. The two strains of Atlantic salmon displayed different seasonal patterns in hypoosmoregulatory ability. The anadromous strain showed a pronounced seasonal pattern with maximal seawater adaptability in early June. In contrast, seawater tolerance in the landlocked strain improved steadily during spring and remained high until late autumn. During the period of enhanced seawater tolerance, hypoosmoregulatory ability increased significantly with body size in both Arctic charr and anadromous Atlantic salmon. The minimum size at which fish were able to regulate plasma sodium following seawater transfer at a level comparable to freshwater levels (<170 mmol I^–1) differed significantly between anadromous Atlantic salmon (ca. 14 cm) and Arctic charr (ca. 22 cm). The results show that seasonal changes in hypoosmoregulatory ability are present in both Atlantic salmon and Arctic charr, and that these physiological traits are retained in the corresponding landlocked strains. However, the seasonal pattern of seawater adaptability as well as the minimum size at which seawater tolerance occurs differs between the two species.     

A test of the hypothesis of pheromone attraction in salmonid migration

Synopsis We tested the hypothesis that anadromous salmonids are guided on their homeward migration by population-specific pheromones. Our findings do not support the hypothesis. Wild migrant Arctic charr,Salvelinus alpinus, from Ikarut River, Labrador were transferred and held in a tributary previously uninhabited by anadromous fish. None of the charr migrating up Ikarut River entered the tributary after fish were transferred. Similarly, migrant charr, which were caught in Ikarut River and released in the tributary below the captive fish, did not remain in the tributary. We re-evaluated the data which have been used to uphold the concept of pheromone attraction in salmonid migration and concluded that support for the hypothesis is unsubstantiated.     

Sacciform cells in the epidermis of the brown trout,Salmo trutta,and the Arctic char,Salvetinus alpinus

Summary Sacciform cells containing an acidophilic, proteinaceous secretion, were identified in the epidermis of the brown trout and Arctic char. This cell type increased in number during the chronic stages of infestation by the ectoparasitic flagellate, Ichthyobodo sp., in immature brown trout, and decreased during sexual maturation in male brown trout and char. It is suggested that the salmonid sacciform cell produces a secretion which protects the fish against infestation or damage by skin parasites.     

Growth hormone and cortisol treatment stimulate seawater tolerance in both anadromous and landlocked Arctic charr

In a comparative experiment the effect of cortisol and growth hormone (GH) on the hypo-osmoregulatory ability of a landlocked and an anadromous strain of Arctic charr (Salvelinus alpinus) was investigated. Cortisol and GH were implanted either alone or in combination, and the fish were exposed to a 24 h seawater challenge test (SWT) on days 14 and 28 after implantation. Hypo-osmoregulatory ability, measured as plasma osmolality and chloride concentration after the SWTs, was better in the anadromous than in the landlocked strain, irrespective of treatment. However, cortisol provided a strong stimulation of hypo-osmoregualtory ability in both strains, and this stimulation seemed to be potentiated by GH in an additive manner. Improved hypo-osmoregulatory ability in GH + cortisol treated anadromous Arctic charr was accompanied by increased gill Na⁺, K⁺-ATPase activity and Na⁺–K⁺–2Cl⁻ cotransporter protein abundance, but no changes in gill Na⁺,K⁺-ATPase α1a and α1b mRNA levels. For landlocked charr the improved hypo-osmoregulatory ability in GH +cortisol treated fish was accompanied only with an increase in gill Na⁺–K⁺–2Cl⁻ cotransporter protein abundance. Hormone treatment caused an improvement of hypo-osmoregulatory ability that was of approximately the same magnitude in the landlocked as in the anadromous Arctic charr. This suggests that the lack of spontaneous development of hypo-osmoregulatory ability often seen in landlocked populations of Arctic charr may depend, at least partly, on a lack of the hormonal activation seen in anadromous populations.     

Spawning behaviour and size-assortative mating of Japanese charr in an artificial lake-inlet stream system

Synopsis We describe the spawning behaviour of Japanese charr,Salvelinus leucomaenis, which have begun to change their life history style from fluvial to lacustrine-fluvial after the construction of an artificial lake by damming. There was a discrete size distribution between mature lake run individuals (larger body size) and mature stream resident fish (smaller body size). Lake run males use pairing tactics, while stream resident males use satellite or pairing tactics depending on the presence of lake run males. Sneaking success of stream resident males is low relative to pairing success of lake run males. Pair formation is strongly assortative by body size and consequently by type. We suggest that both male-male competition and female choice of similar sized males, play an important role in pair formation. We hypothesize that the number of stream resident type relative to lake run type, especially stream resident females, will decrease in future years.     

Ecological segregation within and among Arctic char morphotypesin Gander Lake,Newfoundland

Synopsis Stable isotopes of carbon and nitrogen were used to examine differences in the feeding ecology of sympatric morphotypes of Arctic char, Salvelinus alpinus, from Gander Lake, Newfoundland. Morphotypes were differentiated on the basis of colour (pale and dark) and meristic characteristics. Significant differences in length, age and habitat depth were also noted, with the pale-form being smaller, younger and caught at greater average depths than the dark-form. Pale and dark morphotypes possessed significantly different carbon and nitrogen stable isotope signatures, with the dark-form being more δ¹³C enriched than the pale-form. Within each morphotype grouping distinctive isotopic signature differences related to diet (dark-form) and capture depth (pale-form) were also observed. Differences in the isotopic signatures within and among habitat groupings were induced by differences in feeding opportunities arising from differences in habitat use and feeding strategies and generally corroborated by stomach content analysis and an index of dietary overlap. Study results provide evidence for the contention that sympatric forms of Arctic char in Gander Lake occupy significantly different foraging niches within the lake. Results further suggest ecologically relevant separations within morphotype groupings result in lower resource competition.