Variations in juvenile growth, energy allocation and life-history strategies of two populations of Arctic charr in North Norway

Within the populations of Arctic charr Salvelinus alpinus in Storvatn and Rungavatn, Norway, fish that attained the largest size in the late parr stage migrated as 4+ smolts, fish that attained a medium size became 5+ smolts, and the smallest parr became lake residents. Within the last 2 years of the parr stage, those that became anadromous had a lower growth rate during the winter and early summer than those that became resident, and vice versa in late summer. Thus, anadromous fish grew faster in the parr stage, but in years prior to migration, their growth pattern seemed to differ from that of parr that became resident. Due to early size differences between parr that became anadromous or resident, it is suggested that some of the basis for the decision to smoltify or not in charr depends on the growth rate through the whole parr stage, but with significant modification by the local environment. Rungavatn parr had a significantly higher population density (5 ×), higher lipid content and mortality, an earlier maturation, a lower rate of growth, and only one-third the degree of anadromy than those from Storvatn. It is suggested that the two charr populations have evolved different life-history adaptations due to unequal growth, energy allocation and competition opportunities in the lakes.     

Individual growth and phase differentiation of lacustrine masu salmon, Oncorhynchus masou, under artificial rearing conditions

 In lacustrine masu salmon, Oncorhynchus masou, originated from anadromous fish and inhabiting an artificial lake (Shumarinai Lake), we examined the relationship between individual growth during the juvenile stage and phase differentiation under artificial rearing conditions. In females, the mean fork length of potential (subsequent) 1+ smolts and potential 1+ parr (1+ parr show fish that neither smolted nor matured until 1+ autumn) always differed after their first summer (0+ summer), the former being larger. In males, the juveniles that grew faster during their first spring became 0+ mature male parr. After their first summer, potential 1+ smolts were always larger in body length than potential 1+ mature parr and potential 1+ parr. These results were similar to previous reports of anadromous masu salmon, suggesting that the lacustrine masu salmon studied seems to have maintained the phase differentiation as in the original fish, likely because only 60 years have passed since the formation of the lacustrine population.     

Body composition and energy allocation in life-history stages of brown trout

Energy contents of immature parr and smolts, and mature resident and anadromous brown trout Salmo trutta sampled from a small stream in southern Norway were estimated from lipid, protein and carbohydrate concentrations. In immatures the lipid concentrations were highest in parr in the autumn. Mean lipid concentrations increased significantly with age in parr sampled in autumn (1·3% in age 0+ to 3·4% in age 3+), whereas they did not in smolts. The lipid concentrations of parr in spring were not significantly different from those of similarly aged smolts. By contrast, the relative water content (%) decreased with age in parr in the autumn and increased with age in smolts, mean values being slightly higher in smolts (78%) than in parr (77%). Protein and carbohydrate concentrations did not vary with age in the immature fish, mean protein concentrations being 18·0, 17·5 and 16·8% in parr in the autumn and spring, and in smolts, respectively. In residents, the concentrations of lipids were lower and of water higher, in age group 1 than in older fish, whereas there was no significant variation with age amongst anadromous trout. The energy concentration of 2+ smolts (349 kJ 100 g^-1) was similar to that of 0+ parr in the autumn. Mean somatic energy density in autumn was 1·1 times higher in freshwater residents than in parr at age 1+ (407 and 387 kJ 100 g^-1) and marginally different at age 2+ (462 and 426 kJ 100 g^-1, respectively). The energy contents per unit mass of residents were 1·3–1·6 times that of similar aged smolts. Mean somatic energy density of anadromous trout (504 kJ 100 g^-1) was higher than that of residents (455 kJ 100 g^-1). Somatic energy, lipid and protein concentrations were correlated highly with water contents of all life stages, age and sex groups.     

Genetic architecture of growth and early life‐history transitions in anadromous and derived freshwater populations of steelhead

Heritabilities of growth, precocious maturation and smolting were measured in 75 families of juvenile steelhead or rainbow trout Oncorhynchus mykiss , progeny of within and between line matings (crosses) of wild, anadromous steelhead and wild, resident (lake) rainbow trout originally derived from the same anadromous stock 70 years earlier. The tagged yearling progeny were combined by line in common freshwater rearing containers and graded into three categories: mature, smolt or rearing (undifferentiated) at age 2 years. Heritabilities of precocious male maturity, smolting and growth were moderate to high, and the genetic correlation between growth and smolting was low. Smolting and precocious male maturity were highly variable among families within lines and significantly different between lines. Each of the four lines produced significant numbers of smolts at age two. Smolting and maturation were negatively genetically correlated, which may explain the persistence of smolting in the lake population despite strong selection against lake smolts; balancing selection on male maturation age may help to maintain variation for smolting. The high heritability of smolting, coupled with the inability of smolts that leave the lake to return to it indicates that the genetic potential for smolting can lie dormant or be maintained through a dynamic interaction between smolting and early maturation for decades despite complete selection against the phenotype. The results have significant implications for the preservation of threatened anadromous stocks in fresh water and the inclusion of resident fish of formerly anadromous populations, currently trapped behind long‐standing barriers to migration, as one component of the same population.     

Epidermis thickness and secondary sexual characters in mature male and immature Baltic salmon, Salmo salar L., parr: seasonal variations and effects of castration and androgen treatment

At the breeding season (November) the mature male parr had a thicker epidermis than had the immature parr. In winter (February) there was no difference, while in May, at the smoltification period, the previously mature male parr had a thicker epidermis than the immature parr. In both categories of fish the epidermis was thinner in May than in November. The epidermal thickness in castrated male parr was the same as in immature parr. Treatment with 11-ketoandrostenedione (llkΔ4) increased epidermal thickness in castrated males. Treatment with testosterone and llkΔ4 increased epidermal thickness in previously mature males in spring. There was no development of anadromous secondary sexual characters (hooked lower jaw, adult nuptial colouration) in any male.     

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.     

Mating success of alternative maturation phenotypes in male Atlantic salmon,Salmo salar

Summary Mature male parr successfully fertilized eggs of anadromous female Atlantic salmon, Salmo salar, in competition with anadromous males under simulated natural conditions. Mating situations were created in which mature male parr and anadromous males competing for the opportunity to spawn with an anadromous female differed in allelic forms of the same enzyme. Females deposited eggs into a sequence of 3 or 4 eggs nests. The mean proportion of eggs in a redd fertilized by parr increased with increasing numbers of parr present at a redd, reaching 23% at male parr: anadromous male ratios of 20:1. Single male parr fertilized, on average, 5% of the eggs in a redd. The proportion of eggs in an egg nest fertilized by parr also depended upon the order of egg nest construction, such that parr mating success was highest at the initial nest constructed and lowest at the final nest. Parr have relatively high fertilization success for their size when compared with the smaller maturation phenotype of other salmonids.     

Patterns of Aggression and Operational Sex Ratio Within Alternative Male Phenotypes in Atlantic Salmon

The intensity of male–male aggression during mating is predicted to increase with the operational sex ratio (OSR) (the ratio of sexually active males to females). We observed aggressive behaviour in relation to OSR within alternative phenotypes (large anadromous males and small ‘sneaker’ mature male parr) of Atlantic salmon, Salmo salar, as they competed for access to anadromous females in large outdoor arenas immediately prior to spawning events as well as throughout the spawning season. Anadromous males were primarily aggressive outside of spawning events, and their aggression increased with OSR over a narrow range of 0.08–4. By contrast, mature male parr were aggressive just prior to spawning, with a decrease in aggression over a broad range of OSRs from 1 to 7.4. Similarly, literature data for three other salmonid species indicated a decrease in aggression with increasing OSRs over a range of 1–6.33. These observations suggest that there is considerable variability in competitive behaviour, even within alternative phenotypes. Because our data are derived from repeated observations on the same individuals of different genetic origin, further confirmation of these findings is desirable. Nevertheless, our study underscores the importance of measuring competitive behaviour and OSR at ecologically relevant spatial and temporal scales, both of which may differ between alternative maturation phenotypes.     

Age composition of masu salmon smolts in northern Japan

Smolts of anadromous masu salmon Oncorhynchus masou aged 3+ years were found in a northern Japanese river. This is the first recording of 3+ year smolts in Japan. These fish appeared to originate in the cold upper river where 2+ year parr were found during autumn.     

The Effect of Carlin Tags on Survival and Growth of Anadromous Arctic Charr, Salvelinus Alpinus

Fish tagging methods are important tools in the research and management of salmonids; the Carlin tag is widely used, especially in anadromous salmonids. However, this tagging method may interfere with fish behaviour and increase post-smolt mortality. To estimate Carlin-tag-inflicted mortality in Arctic charr, we used jet injection of alcian blue (colour tags) as a control. Recapture rates were higher for the colour-tagged fish than for the Carlin-tagged fish for both one- and two-year-old smolts. For one-year-old smolts in 1992, the recapture rate was 18.9% for colour-tagged fish and 10.0% for Carlin-tagged ones. For two-year-old smolts in the same year, 34.0% of colour- and 28.6% of Carlin-tagged fish were recaptured; in 1993, recapture rates were 17.2% and 6.3% for colour- and Carlin-tagged fish, respectively. Colour-tagged one-year-old Arctic charr also grew faster than their Carlin-tagged counterparts. No differences in growth rates were found between tagging methods in two-year-olds. These results imply that for hatchery-reared Arctic charr, survival rates using Carlin tagging may be half to one third that of colour tagging. Growth was also higher among colour-tagged fish, indicating that Carlin tags alter the behaviour of fish in the sea.     

Growth and development of a bimodal length-frequency distribution during smoking in a wild population of white-spotted charr in northern Japan

Smolt age and size, and individual growth during smoking were examined in a wild population of white-spotted charr, Salvelinus leucomaenis , in a small stream in northern Japan. Smolt size (110–210 mm F.L.) differed neither between sexes nor among age cohorts of 2–7 years. Back-calculation of riverine growth of smolts using otoliths showed a gradually increasing discrepancy in fork lengths of these age cohorts in both males and females, the faster-growing individuals smolting at an earlier age. A mark—recapture study on individually-identified fish revealed that the initial (spring-summer) length-frequency distribution of parr was unimodal, but became slightly skewed to the larger side in autumn, resulting in a bimodal distribution being evident by the following spring, with the upper modal group primarily composed of smolts. However, closer examination revealed that both male and female potential (subsequent) smolts were larger than the potential parr in the previous spring, the minimum lengths of the former being around 90–100 mm f.l. Specific growth rates of the potential smolts remained relatively high after summer through to the following spring, whereas those of the potential parr decreased after summer, despite having been similar between the two groups during the previous spring and summer. These observations revealed that the developmental triggering of smelting possibly occurs in or following the summer season.     

The timing, sex and age composition of the wild and reared Atlantic salmon ascending the Simojoki River, northern Finland

In the Simojoki River in the northern Gulf of Bothnia, reared salmon stocked as smolts produced considerable numbers of ascending one‐sea‐winter (1 SW) males, whereas the proportion of male 1 SW salmon was low among spawning migrants of wild or reared parr origin. The sex ratio among ascending wild fish and reared salmon stocked as parr was similar, with females predominating, while reared salmon stocked as smolts were mainly males. The multi‐sea‐winter (MSW) salmon entered the river annually within a fairly short time period from the beginning of the migration season, independent of their sex or origin. 1 SW males migrated into the river significantly later in the season than MSW males. The results indicate that the delayed opening of the fishing season in the Gulf of Bothnia is effective in reducing the harvest of MSW salmon at sea. However, as the timing of the ascent may vary by several weeks from year to year, the effect of this regulation bound to certain calendar days may also vary considerably from year to year.     

Mating of anadromous Atlantic salmon, Salmo salar L., with mature male parr

Sexually mature male parr successfully mated with female anadromous Atlantic salmon, Salmo salar , in the absence of anadromous males. There was no significant difference between the proportion of eggs fertilized by mature male parr and anadromous males. One of the demographic consequences of our observations is that overfishing could eventually eliminate anadromy in male members of a population.     

Development of length–bimodality and smolting in wild stocks of Atlantic salmon, Salmo salar L., under different growth conditions

Growth dynamics of juvenile Atlantic salmon, Salmo salar L., from two sections of the Narcea River and one of the Esva River (Northern Spain) were examined in relation to the development of bimodality in their size–frequency distributions. Size–bimodality was clearer under intermediate growth (section A) than under relatively fast or slow growth. The proportion of fish entering the upper modal group increase with growth intensity. Composition of upper and lower modal groups became fixed prior to December, and at this time both groups separated on the 90–95 mm interval. Fish exhibiting smolt appearance in late March (larger than 130 mm) had already been upper group fish in December, while parr-like fish and those that remained in the river by May (potential 2-year-old smolts) had formed the lower modal group. Anadromous salmon catch in the Narcea River was mostly of previously 1-year-old smolts (97.6%), of which 94% were larger than 100 mm by their first winter. In the Esva River, slow growth of juveniles is consistent with a large proportion of 2-year-old smolts (47.9%) among anadromous salmon. Both juvenile samples and scale analysis of anadromous salmon indicate that 2-year-old smolts were larger than 1-year-olds. Early disappearance of the former (before March) is, at least, partially related to earlier migration of large fish, since sexual maturity of parr does not provide a complete explanation. The Narcea stock have a minimum length at smolting of about 130 mm and an optimum smolt size in the 155–175 mm interval. Mean smolt length did not vary although the winter length changed between years.     

Relationship between metabolism, sex and reproductive tactics in young Atlantic salmon (Salmo salar L.)

Atlantic salmon can differ markedly in their growth and in the timing of reproductive maturation, leading to the dramatic contrast between the large anadromous adults and the diminutive mature male parr. This study examined the growth rates, anatomical and physiological characteristics of parr during the adoption of their discrete life histories to ascertain whether these properties can explain tactic choice. To minimise the impact of habitat differences upon these attributes, salmon were reared in the laboratory until 1.5years of age, when the "decisions" to undergo smoltification or to mature as parr had been taken. At 1.5years, both males and females showed bimodal size-frequency distributions. Neither the population of origin nor the paternal reproductive tactic influenced the "decision" to mature or the growth trajectories. Growth rate (% massday(-1) during their final 10months) and the % male and female offspring in the upper modal group were strongly correlated and varied markedly among families. Mean growth rate per family was negatively correlated with mean metabolic rate per family at emergence. Growth rate decreased as a function of parr size in January and the growth rates of upper modal fish were displaced upwards relative to those of lower modal fish. Most males in the smaller size mode matured, whereas all other fish began smoltification. Mature male parr did not differ from similarly sized female pre-smolt in routine metabolic rate, but these smaller fish had higher metabolic rates than larger male and female pre-smolts. However, mature parr differed markedly from similarly sized females and from larger male and female pre-smolts in possessing higher oxidative and lower glycolytic capacities in muscle. Overall, these data are consistent with the interpretation that growth rates dictate the distribution of parr between upper and lower modal groups. Individuals from faster growing families would be more likely to pass the threshold for smoltification and to accelerate growth, whereas those from slower growing families would remain in the lower mode. The use of metabolic capacities, e.g. metabolic rate, was linked with modal group, whereas muscle oxidative capacity was linked with male maturity. Mean family metabolic rate at emergence was negatively linked with mean growth during the subsequent year, suggesting that metabolic efficiency facilitates growth and eventually smoltification.     

Smolting characters in anadromous and resident Arctic charr, Salvelinus alpinus (L.)

Morphological differences, haematocrit value and chloride cells were examined in downstream migrating Arctic charr, Salvelinus alpinus (L.) from the Hals River, North Norway, and resident charr from Lake Storvann. Fish were classified as visual parr, silvery parr or smolt based on the degree of silvering and the Occurrence of lateral parr marks. On average, 47% of downstream migrating Arctic charr were classified as visual molts, but only 14% of the resident charr. Charr longer than 20 cm fork length were mainly classified as visual molts. Morphometrical analyses of body size and shape revealed that most of the variations could be explained in terms of variations in fork length. Length adjusted ratios of post-anal distances were significantly higher in migrating charr than in resident charr. The number of developed chloride cells, the cell nucleus diameter in the basin of secondary gill lamellae and blood haematocrit values were significantly higher among downstream migrating charr than in resident charr. However, morphological differences between anadromous and resident Arctic charr were not necessarily synchronized with the development of chloride cells.     

Does the Refuge Availability Influence the Spawning Behavior of Mature Male Parr in Salmonids? A Test in the Miyabe Charr

In many salmon and charr, both 'migratory males' and 'precocious males' are maintained as alternative life-histories within a population. During breeding, precocious males gather around the mating pair of migratory fish and attempt to sneak in the spawning between larger male and female charr. Refuges such as debris and shallow areas may provide shelter from aggressive attacks and allow precocious males to hold positions closer to the pair (nest) and/or sneak more easily during the mating. Here we examine whether the availability of refuges influences the breeding behavior of precocious male parr of the Miyabe charr under experimental and natural conditions. Aggressive attacks directed toward precocious parr by the mating pair or other parr was not influenced by the refuge availability either under experimental conditions or in the wild. We did not find the effect of high refuge availability on the proximity of parr to the nest beyond the experimental conditions where the density of refuges was probably higher than in the natural stream. There were no effects of high refuge availability either on parr's attempts at sneak mating or on their success in spawning. Thus, our results provide little evidence that the refuge availability influences parr sneaking behavior. Other possible effects of high refuge availability, for example, the increase of fertilization success and the decrease of energy expenditure and risk taking, remain to be considered.     

Relationships between Na+, K+-ATPase, sialic acid content and fluidity in rainbow trout density-separated erythrocytes

Smolt of anadromous Arctic charr Sahelinus alpinus (L.) migrating from Storvatn, northern Norway, had a similar feeding intensity as resident Arctic charr from the benthic- and pelagic zone in early summer in fresh water. While smolts fed mostly on surface insects, however, resident Arctic charr had a more broader diet consisting mostly of chironomids, Bosmina sp., benthic prey and surface insects. This indicates that Arctic charr smolts have adopted a surface oriented feeding behaviour prior to their sea migration.     

Paternal reproductive strategy influences metabolic capacities and muscle development of Atlantic salmon (Salmo salar L.) embryos

Male Atlantic salmon follow a conditional strategy, becoming either "combatants" that undertake a seaward migration and spend at least a year at sea or "sneakers" that remain in freshwater and mature as parr. A variety of physiological indices showed significant but small differences between the offspring of males that use these two reproductive tactics. Offspring fathered by anadromous male Atlantic salmon (Salmo salar L.) showed greater muscular development and muscle metabolic capacities but lower spontaneous movements than those fathered by mature male parr. At hatch and at maximum attainable wet weight (MAWW), offspring fathered by anadromous males had higher activities of mitochondrial (cytochrome C oxidase and citrate synthase) and glycolytic (lactate dehydrogenase [LDH]) enzymes than progeny of mature male parr. Enzymatic profiles of progeny of anadromous fathers also suggested greater nitrogen excretion capacity (glutamate dehydrogenase) and increased muscular development (creatine kinase and LDH) than in the progeny of mature parr. At MAWW, juveniles fathered by mature parr made considerably more spontaneous movements, presumably increasing their energy expenditures. For juveniles fathered by anadromous males, total cross-sectional areas of white and red muscle at hatch were higher due to the greater number of large-diameter fibers. We suggest that the slightly lower metabolic capacities and muscular development of alevins fathered by mature parr could reflect differences in energy partitioning during their dependence on vitellus. Greater spontaneous movements of offspring of mature male parr could favor feeding and growth after the resorption of the vitellus.     

Divergence in smolt production from the stocking of 1-summer-old and 1-year-old Atlantic salmon parr in a northern Baltic river

One‐year‐old and 1‐summer‐old salmon parr have been stocked in the Simojoki, a northern Baltic river, to protect the natural salmon stock from extinction. This enhancement practice was studied during a 5‐year stocking period, when an average of 14.3 1‐summer‐old parr and 5.4 1‐year‐old parr was needed to produce one sea running smolt. The mean yield from stocking was 70 smolts per 1000 stocked 1‐summer‐old parr and 186 smolts per 1000 stocked 1‐year‐old parr. The mean cost of smolts produced by stocking 1‐year‐old and 1‐summer‐old parr was 1.2 and 2.6 times the cost of producing one 50 g smolt in the hatchery, respectively. There was a negative relationship between density of wild parr and length of smolts stocked as 1‐summer‐old parr in the river, but no relationship between wild parr density and length of smolts stocked as 1‐year‐old parr. This indicated that 1‐summer‐old parr were more susceptible than the older parr to competition with wild parr. Considering production costs and smolt yield, stocking of 1‐year‐old parr appeared to be more profitable than that of 1‐summer‐old parr in the enhancement of the endangered salmon stock.