Effects of food availability on temporal activity patterns and growth of Atlantic salmon

1. Patterns of sheltering and activity are of fundamental importance in the ecology of animals and in determining interactions among predators and prey. Balancing decreased mortality risk when sheltering with increased feeding rate when exposed is believed to be a key determinant of diel patterns of sheltering in many animals. 2. Despite lower foraging efficiency at night than during the day, Atlantic salmon Salmo salar parr are nocturnal during winter and at low summer temperatures. Nocturnal activity also occurs at warm water temperatures during summer, but little is known about the functional significance of this behaviour. 3. This study aimed to determine: (1) the preferred activity and shelter pattern of Atlantic salmon parr during warm summer months, and (2) their response to variations in food availability when balancing growth rate (G) and mortality risk (M), as expressed through time out of shelter. We differentiated among four potential responses to reduced food availability: (1) no response; (2) G decreases but M remains constant; (3) G remains constant but M increases; and (4) G decreases and M increases. 4. Time exposed from shelter was inversely related to food availability. Fish subject to high food availability were significantly less active during the day than those with restricted rations. However, food availability had no significant effect on the extent to which fish were active at night. There was no evidence of variation in growth rate with food availability. 5. Salmon were predominantly nocturnal at high ration levels, consistent with their previously reported behaviour during winter. Rather than switching to diurnal behaviour at high temperatures per se, as previously was supposed, it appears that the fish are diurnal only to the extent needed to sustain a growth rate, and this extent depends on food availability. 6. Atlantic salmon parr modulate the amount of time they are active rather than growth when responding to variations in food availability over an order of magnitude.     

Effect of actinomycin-D on gill Na+-K+ ATPase of juvenile chinook salmon, Oncorhynchus tshawytscha (Walbaum)

Actinomycin-D reduced gill Na⁺-K⁺ ATPase activity of chinook salmon Oncorhynchus ishawytswha (Walbaum), smolts and saltwater-adapted juveniles but had no significant effect on the enzyme activity of parr in fresh water. The similarity in response suggests that even though smolts are found in fresh water, their enzyme system is more characteristic of saltwater-adapted juveniles than freshwater-dwelling parr. A greater reduction in enzyme activity was found when fish were treated with actinomycin-D in salt water than in fresh water, suggesting that the enzyme degradation rate wasgreater in salt water than in fresh water.     

Seasonal variations in the prevalence and infestation intensity of Gyrodactylus salaris Malmberg, 1957 (Monogenea: Gyrodactylidae) on Atlantic salmon parr, Salmo salar L., in the River Batnfjordselva, Norway

The prevalence of Gyrodactylus suluris Malmberg. 1957 on both yearlings and older parr of Atlantic salmon ( Sulmo sulur L.) in the River Batnfjordselva was 100% through most of the year. With one exception, uninfested fish were only found in the winter and spring after the water temperature had fallen to almost 0° C for 2–3 months. In general, the abundance (i.e. the mean number of parasites per investigated fish) of G. salaris increased during the warm period of the year (summer and early autumn). Abundanceas high as 1153 and 4418 in early autumn was found on yearlings and older parr, respectively. The abundance decreased during the cold period of the year (winter and early spring), in some cases to as low as two and four G. salaris on yearlings and older parr, respectively. About 86% of the G. salaris specimens were found on the fins of the salmon parr: mainly on the dorsal fin (34.4%) and the pectoral fins (27.0%). The remainder of the parasites were distributed on the body (7.8%), the head (3.5%), and the gills (2.6%).     

Interhabitat and instream movements of large Atlantic salmon parr in a Newfoundland watershed in winter

Radio‐telemetry was used to investigate movement of large, mainly mature male (80%) Atlantic salmon Salmo salar parr in Stoney River, Newfoundland during early winter (November; water temperature 6·0 ± 0·1° C) and mid‐winter (January to February; 0·8 ± 0·0° C). Site fidelity of parr in early winter was low. Parr moved between fluvial and lacustrine habitats and were active throughout the diel cycle. Parr caught in fluvial habitats in mid‐winter were smaller and younger than parr caught in early winter. Site fidelity of parr in mid‐winter was greater than in early winter. Parr in mid‐winter moved between fluvial and adjacent small lacustrine habitats, but avoided a larger pond inhabited by large piscivorous fishes. Instream movement rates in mid‐winter were lower than in early winter and occurred primarily during hours of darkness (dawn, dusk and night). Fluvial habitats were relatively stable and ice‐free throughout the study periods. These results suggested that large Atlantic salmon parr utilize a variety of habitats and remain active throughout the winter, even under stable environmental conditions.     

Use of space and food by resident and migrant brown trout,Salmo trutta

Synopsis Parr and resident forms of brown trout,Salmo trutta, from Vangsvatnet Lake, Norway live in freshwater, while migrant forms live in coastal waters during summer and in freshwater during winter. About 80% of parr and residents live at depths <5 m, smolts and migrants are more confined to near-surface water. Brown trout partly segregate by size, age and sex from spring through autumn. More than 90% of parr and residents in the tributaries are 0–2 years old, 2–14 cm in length, in the littoral zone 0–3 years old, 7–24 cm in length, and in the pelagic zone 2–6 years old, 18–32 cm in length. The mean body length of equal-aged fish increases from tributaries through littoral to pelagic zones in the lake. Smolts (2–7 years, 14–29 cm) leave the lake from April through August and return during September–October. Migrants (2–11 years, 23–67 cm) leave the take in April–May and return during August–September; sexually mature fish return earlier than immatures. Female brown trout are less stream-dwelling, but more migrant and pelagic than males. Most individuals in the lake population spend the winter in the littoral zone. In the tributaries, diet differs significantly between age-groups of parr; young fish feed on smaller food items than do older fish. In the lake, parr and residents living in the same habitats feed on the same food items. Littoral brown trout feed mainly on insect larvae and chironomid pupae, pelagic brown trout feed on zooplankton and surface insects. Migrants feed little while staying in freshwater, except for matures which feed on young salmonids and surface arthropods during the 2 first months after they had returned from coastal waters. The results are discussed in relation to growth possibilities and mortality risks of the different habitats.Reprint request to B. Jonsson.     

Influence of size, winter duration and density on sexual maturation of Atlantic salmon (Salmo salar) juveniles in Little Codroy River (southwest Newfoundland)

Factors affecting sexual maturation of Atlantic salmon ( Salmo salar L.) juveniles in fresh water were analysed under wild conditions in Little Codroy River (southwest Newfoundland). Large size after the first year of life and short winters favoured maturation of age 1 + male parr. Incidence of maturing 1+ males was greater where high densities of parr (>0.2 individuals m ²; fish >6cm fork length) were recorded in the second season of growth. These results were discussed according to current knowledge on early maturation in Atlantic salmon.     

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.     

Choice of feeding station in Atlantic salmon, Salmo salar, parr: effects of predation risk, season and life history strategy

The current speed at which underyearling salmon parr held feeding stations was examined from late summer to early winter in laboratory flume tanks that offered a choice between (a) areas with high water flow, high food availability but high predation risk and (b) areas with low flow, little food but shelter from predators. In August, those fish that would become smolts aged 1 + (and which by late winter formed the upper modal group, UMG, of the bimodal size distribution) adopted positions in faster currents than did the fish which would take a further year to reach the smolt stage (the lower modal group, LMG). However, the chosen current speed of UMG fish decreased through the period of study, so that by December all fish were found in areas of low flow, and hence little food. Both date and water temperature had independent effects on the chosen current speed of UMG fish.
The effect of predation risk was investigated using a model trout. A brief sight of this predator caused 47% of fish that had been in the main, exposed currents to move to slacker, sheltered areas; they took 1 h, on average, to return to their previous position. The fish that remained in position upon seeing the predator reduced their rate of tail beating, presumably increasing crypticity. Eventual UMG parr were less likely than were LMG fish to move away upon seeing a predator. The fish moved to faster currents than normal 2–3 h after seeing the predator, possibly compensating for the earlier reduction in feeding rate.     

Life histories of Atlantic salmon altered by winter temperature and summer rearing in fresh- or sea-water

Increased growth during winter increased the incidence of age 1+ Salmo salar smolts in spring. High condition factor in spring and good growth in summer was associated with a high incidence of sexually mature males in autumn. In two experiments, groups (n=160–237 per group) of individually identified parr, either ungraded (lower and upper modal groups: LMG, UMG) or size-graded (LMG only), were reared at either 10, 6 or 3 °C overwinter (Nov to May). At 10 °C, up to 51% of parr originally in the LMG became smolts in spring at age 1+. In contrast, at either 6 or 3 °C (control), < 6% of LMG parr became smolts. The probability of being recruited into the UMG overwinter was positively related to initial body size, and was increased by size-grading. Smolt recruitment was two-fold higher among females compared to males; a proportion of males by age 0+ had already opted to mature at age 1+ rather than smolt at age 1+. In contrast, smolting at age 1+ was not inhibited in males previously mature at age 0+. During summer (May to Nov), all experimental groups were reared at ambient temperature, each subdivided between fresh water (max 21 °C) or seawater (max 15 °C). Good growth in seawater of winter recruits into the UMG confirmed they had completed smolting. Mortality in seawater among parr was 41–83%, and among smolts was 10–22%. Specific growth rate during summer was inversely related to winter rearing temperature. The incidence of sexual maturity in autumn at age 1+ among male parr was positively related to winter rearing temperature, fork length and condition factor in May, but there was large variation among individuals with respect to body size and maturity. Summer rearing in seawater reduced growth and the incidence of maturation. Parr and post-smolt maturity was 84–99% and 100% in fresh water respectively, 21–58% and 0% in seawater.     

Effect of stripping on the length, condition and gonadal state of mature male Atlantic salmon, Salmo salar L., parr during autumn and winter

Hatchery-reared mature male Atlantic salmon parr were shorter than immature parr of the same age through the winter period and this difference increased prior to smolting in spring. Stripping had no effect on growth. In autumn mature male parr had a higher condition factor (CF) than immature parr, but from early January no difference was observed between unstripped mature males and immature parr. Stripping reduced the CF of male parr but the difference had disappeared by the time of smolting when the CFs of both mature groups were lower than that of immature fish. The quantity of expressible milt in stripped males decreased from c . 3% body weight in November and December to <0.05% by 2 March, after which no further milt expression was recorded. Milt expression in previously unstripped males was about 1% of body weight on 2 March, similar to that of stripped males 1 month earlier.     

Redistribution of juvenile salmonid fishes after localized catastrophic depletion

Juvenile Atlantic salmon and brown trout were depleted at three sites ( c . 108–380 m²) of a natural stream during the summer months of 1991 and 1992. Local population changes and movements of fish marked in sections adjacent to each depleted area were monitored thereafter. There was very little movement of marked salmon parr into the central regions of the depleted areas following the immediate post-marking period. Upstream movement by young-of-the-year fish from high density sections in mid-late summer was noted for trout but not salmon. Unmarked 1-year-old salmon parr immigrated into depleted areas in June 1992, and the pattern of recolonization was consistent with migration upstream from the adjoining river. It is concluded that resident salmon were very strongly site-attached and resource tracking was of no functional significance as a compensatory mortality mechanism. The occurrence of a long distance migratory component in the population during early-mid summer indicates that this, rather than local resource tracking, constitutes a potential compensatory mechanism.     

Ice‐dependent winter survival of juvenile Atlantic salmon

Changes in snow and ice conditions are some of the most distinctive impacts of global warming in cold temperate and Arctic regions, altering the environment during a critical period for survival for most animals. Laboratories studies have suggested that reduced ice cover may reduce the survival of stream dwelling fishes in Northern environments. This, however, has not been empirically investigated in natural populations in large rivers. Here, we examine how the winter survival of juvenile Atlantic salmon in a large natural river, the River Alta (Norway, 70°N), is affected by the presence or absence of surface ice. Apparent survival rates for size classes corresponding to parr and presmolts were estimated using capture‐mark‐recapture and Cormack‐Jolly‐Seber models for an ice‐covered and an ice‐free site. Apparent survival (Φ) in the ice‐covered site was greater than in the ice‐free site, but did not depend on size class (0.64 for both parr and presmolt). In contrast, apparent survival in the ice‐free site was lower for larger individuals (0.33) than smaller individuals (0.45). The over‐winter decline in storage energy was greater for the ice‐free site than the ice‐covered site, suggesting that environmental conditions in the ice‐free site caused a strong depletion in energy reserves likely affecting survival. Our findings highlight the importance of surface ice for the winter survival of juvenile fish, thus, underpinning that climate change, by reducing ice cover, may have a negative effect on the survival of fish adapted to ice‐covered habitats during winter.     

Comparative diets and foraging strategies of subyearling Atlantic salmon,brown trout,and rainbow trout during winter

Over‐winter survival of salmonids in streams is thought to be an important population regulation mechanism. Yet because of the difficulty of conducting field studies due to adverse weather or ice conditions, compared to other seasons, salmonid ecology during winter is least understood. Consequently, we sought to examine interspecific feeding associations of an important salmonid stream assemblage in the Lake Ontario watershed during winter. The diets of Atlantic salmon (Salmo salar) parr, brown trout (S. trutta) parr, and rainbow trout (Oncorhynchus mykiss) parr were significantly different in February but not in March. Salmonid diets differed from the benthos and the drift during both months. Dipterans (chironomids, simuliids, and tipulids) and ephemerellids were the major prey taxa consumed. All three species fed more heavily on prey items from the benthos than from the drift. The diet of Atlantic salmon had the highest similarity to the benthos whereas the diet of brown trout had the lowest similarity to the drift. All three salmonid species generally selected ephemerellids, limnephilids, and chironomids and avoided elmids. These winter feeding observations are the first reported for this specific salmonid assemblage and will help managers better understand interspecific associations during this critical period.     

Seasonal cycle in vitamin A1/A2-based visual pigment composition during the life history of coho salmon (Oncorhynchus kisutch)

Microspectrophotometry of rod photoreceptors was used to follow variations in visual pigment vitamin A₁/A₂ ratio at various life history stages in coho salmon. Coho parr shifted their A₁/A₂ ratio seasonally with A₂ increasing during winter and decreasing in summer. The cyclical pattern was statistically examined by a least-squares cosine model, fit to the 12-month data sets collected from different populations. A₁/A₂ ratio varied with temperature and day length. In 1+ (>12 month old) parr the A₂ to A₁ shift in spring coincided with smoltification, a metamorphic transition preceding seaward migration in salmonids. The coincidence of the shift from A₂ to A₁ with both the spring increase in temperature and day length, and with the timing of seaward migration presented a challenge for interpretation. Our data show a shift in A₁/A₂ ratio correlated with season, in both 0+ (<12 months old) coho parr that remained in fresh water for another year and in oceanic juvenile coho. These findings support the hypothesis that the A₁/A₂ pigment pair system in coho is an adaptation to seasonal variations in environmental variables rather than to a change associated with migration or metamorphosis.     

Effect of habitat types on survival, spatial distribution and production of an allopatric cohort of Atlantic salmon, Salmo salar L., under conditions of low competition

An allopatric cohort of Atlantic salmon, Salmo salar L., introduced to a small previously fishless stream was studied from parr to the smolt stages. In May 3900 0+ parr (mean total length 30mm) were planted at three different densities in habitats with slow, intermediate and fast water velocities. During the first year, high mortality occurred during the first 7 weeks after planting in May, and in September–October. Survival from May 1985 to April 1986, before the smolt emigration, was24.8%. The smolt yield 1 year after planting was 15.5%. It is suggested that the high survival was caused by low competition. Most of the redistribution of the fish took place during the first months. Type of planting habitat affected the timing of redistribution. The parr left slow-flowing, deep habitats with fine substrate soon after planting, while redistribution was slowest in the fastest flowing habitats with coarse substrate. The observed avoidance of slow, deep habitat types in the absence of interspecific competition, suggests that this may be a fixed behavioural response, and not due to competition. Long movements, up to 800 m, were recorded only within the first 7 weeks after planting. The effect of planting densities on population density was most pronounced immediately after planting in the fast and also intermediate habitats. Planting density effects declined and were not detectable after 1 year. The effect of habitat type on fish numbers and biomass was pronounced irrespective of planting densities. Growth was fastest in the intermediate habitat, and at the lower planting densities. Production was 7.2 g m⁻² the first summer-autumn. Due to smolt emigration, few fish remained in the stream the second summer-autumn, and the production was 1.0 g m⁻².     

Neutrophils and B-cells in blood and head kidney of Atlantic salmon (Salmo salar L.) challenged with infectious pancreatic necrosis virus (IPNV)

Salmon B-cells and neutrophils were studied by flow cytometry in IPNV infected salmon. A highly virulent strain of IPNV was used for challenge of parr and post-smolts. The parr were challenged by intraperitoneal (ip) injection while salmon post-smolts were challenged by ip injection or cohabitation. No mortality occurred in the parr groups, but a cumulative mortality of about 50% was obtained in cohabitant infected post-smolt groups and less than 10% in ip challenged post-smolts. The virus levels were low in head kidney (HK) samples from survivors compared to dead fish. The percentages of neutrophilic granulocytes and Ig+ cells (B-cells) were analysed using HK and blood samples from survivors. The cell populations were identified by monoclonal antibodies (MAb) E3D9, recognising neutrophils, and G2H3 recognising Ig+ cells (B-cells). Parr sampling for leucocyte analyses took place about 1.5 weeks prior to and about 4 weeks post challenge. This corresponded to about 8 and 2.5 weeks before the fish were adapted to seawater transfer. In parr head kidney leucocytes (HKL) we observed significantly lower (p < 0.05) levels of neutrophils in ip infected fish compared to non-infected control fish. The post-smolt sampling from infected fish took place 2 weeks prior to and in the fifth and sixth week post challenge. HKL samples from both surviving cohabitants and ip injected fish had significantly (p < 0.05) lower levels of neutrophils than non-infected control fish. The cohabitant fish also had significantly (p < 0.05) higher levels of B-cells in HKL compared to ip injected fish. No significant changes in B-cells in HKL or peripheral blood leucocytes (PBL) was observed in infected parr or ip infected post-smolts compared to control fish. The relative leucocyte levels of the fish prior to challenge and in non-infected control fish are in accordance with earlier findings. The results indicate that non-specific immune cells like neutrophils are highly influenced by IPNV infection of parr and post-smolts several weeks post challenge.     

Flavobacterium psychrophilum associated with septicaemia and necrotic myositis in Atlantic salmon Salmo salar: a case report

We describe the first case from Norway of increased mortality in Atlantic salmon Salmo salar (L.), with septicaemia and necrotic myositis, associated with infection by Flavobacterium psychrophilum. The outbreak occurred in smolt of 60 to 100 g in fresh water on a land-based farm in Western Norway during winter 2008-2009. The water temperature was < 5 degrees C and the accumulated mortality was 7.0%. Necropsy of dead and moribund fish revealed a swollen dark spleen, pale liver, serohaemorrhagic ascites and haemorrhage in the abdominal fat and muscle. F. psychrophilum was isolated from the kidney and spleen of diseased fish. Muscle biopsy revealed the presence of long filamentous rods in necrotic areas of skeletal muscle. Immunohistochemistry was positive for F. psychrophilum. Identification of cultured isolates as F. psychrophilum was confirmed using phenotypic testing and sequencing of the 16S rRNA gene. Analysis by allele-specific polymerase chain reaction (allele-specific PCR) indicated that 2 different genotypes of the bacterium were present in the outbreak.     

Interactions between riparian shading and food supply: a seasonal comparison of effects on time budgets, space use and growth in Atlantic salmon Salmo salar

This study examines seasonal (winter v. summer) differences in space-time budgets, food intake and growth of Atlantic salmon Salmo salar parr in a controlled, large-scale stream environment, to examine the direction and magnitude of shifts in behaviour patterns as influenced by the availability of overhead cover and food supply. Salmo salar parr tested in the presence of overhead cover were significantly more nocturnal and occupied more peripheral positions than those tested in the absence of overhead cover. This increase in nocturnal activity was driven primarily by increased activity at night, accompanied by a reduction in daytime activity during winter. The presence of overhead cover had no effect on rates of food intake or growth for a given food supply in a given season. Growth rates were significantly higher for fish subjected to a high food supply than those subjected to a low food supply. Food supply did not affect the extent to which S. salar parr were nocturnal. These results were consistent between winter and summer. The use of riparian shading as a management technique to mitigate the effects of warming allows the adoption of more risk-averse foraging behaviour and may be particularly beneficial in circumstances where it serves also to increase the availability of food.     

Salinity tolerance of the mosquito fish, Gambusia affinis (Baird and Girard)

Experiments were conducted to determine the adaptability of mosquito fish to various salt concentrations. Mosquito fish, Gambusia affinis , (total length (T.L.) 27–37 mm) were subjected to abrupt and gradual changes from fresh water to various salinities (S). Low mortality (10%) occurred when the fish were transferred from fresh water (S = 0·5‰) to 50% sea water (S = 19·5‰ Through gradual adaptation 65·0% of the fish were able to tolerate 100% sea water for 7 days. After a 7 -day stay in sea water, fish were readapted to fresh water during a 3-h period. The results indicate that they were well able to tolerate the abrupt change. Through a gradual adaptation fish were also able to tolerate salinities ranging from 39·0‰ (100% sea water) to 58·8‰     

Movements and habitat use by PIT-tagged Atlantic salmon parr in early winter: the influence of anchor ice

1. Movements and habitat use by Atlantic salmon parr in Catamaran Brook, New Brunswick, were studied using Passive Integrated Transponder technology. The fish were tagged in the summer of 1999, and a portable reading system was used to collect data on individual positions within a riffle‐pool sequence in the early winter of 1999. Two major freezing events occurred on November 11–12 (Ice 1) and November 18–19 (Ice 2) that generated significant accumulations of anchor ice in the riffle. 2. Individually tagged parr (fork length 8.4–12.6 cm, n = 15) were tracked from 8 to 24 November 1999. Over this period, emigration (40%) was higher from the pool than from the riffle. Of the nine parr that were consistently located, seven parr moved <5 m up‐ or downstream, and two parr moved more than 10 m (maximum 23 m). Parr moved significantly more by night than by day, and diel habitat shifts were more pronounced in the pool with some of the fish moving closer to the bank at night. 3. During Ice 2, there was relatively little movement by most of the parr in the riffle beneath anchor ice up to 10 cm in thickness. Water temperature was 0.16 °C above the freezing point beneath anchor ice, suggesting the existence of suitable habitats where salmon parr can avoid supercooling conditions and where they can have access to low velocity shelters. To our knowledge, these are the first data on habitat use by Atlantic salmon parr under anchor ice.