Winter behaviour of juvenile Atlantic salmon Salmo salar L. in experimental stream channels: effect of substratum size and full ice cover on spatial distribution and activity pattern

Activity and choice of areas offering different cover (substratum or surface ice) for juvenile Atlantic salmon Salmo salar were studied in experimental stream channels during winter. Channels were completely ice covered between December and March. During this period, the ice thickness increased from 50 to 300 mm after which 50% of the ice was experimentally removed and followed by c. 2·5-fold increase in discharge to simulate the effects of spring flood. Large substrata provided preferred habitats but areas with small substratum sizes were also used when full surface ice provided above-stream cover and the stream discharge was relatively low. The fish remained nocturnal throughout the study but the level of day activity significantly increased as the surface ice became thicker. Maximum movement distance during a 24 h period and homing-at-dawn behaviour remained at a constant level throughout the main winter, but significantly changed during the simulated spring flood (mean ± s . e . maximum extent of movements within 24 h increased from 1·1 ± 0·1 to 3·0 ± 0·5 m; homing behaviour decreased from the highest level of 89·3 to 34·6% during spring flood). Overwinter survival was high (92·9%). Relative mass increase during the study ranged from –8·3 to 28·5%, and 84% of the juvenile Atlantic salmon gained mass. The highest rates of mass increase were associated with frequent movements between areas of different substratum size. The results indicate that during winter: (1) Atlantic salmon parr preferred large substratum cover compared with surface ice cover at the fish densities studied here, (2) juvenile Atlantic salmon were predominantly nocturnal but diurnal activity increased as surface ice became thicker and (3) increase in water discharge during spring altered the behaviour of juvenile Atlantic salmon and may have caused additional habitat shifts.     

Maturation of male age-0 Atlantic salmon following a massive, localized flood

Maturation of male age-0 Atlantic salmon Salmo salar parr in New England, U.S.A. streams is rare (˜5%), but age-0 parr maturation was high (74%) by autumn in the Sawmill River following a massive, localized flood. Maturation was low in two other study streams (3, 7%) in the same year as the flood, and in the Sawmill River (6%) and the other rivers (5%) in the subsequent year, suggesting that high maturation rates were related to the flood. The high age-0 maturation rates appear to have been the result of greater growth opportunity following the flood. Masses of fish in October were two-fold greater in the Sawmill River (13·2 g) than in the other rivers (6·5, 6·9 g). Mechanisms contributing to the fast growth may include community reorganization following the flood and water temperature differences among rivers. The flood caused an age-0 year-class failure for brook trout Salvelinus fontinalis and brown trout Salmo trutta and a large reduction (69%) in the number of salmon compared to the other rivers, possibly reducing competition or agonistic interactions among remaining fish. Average water temperatures were slightly warmer in the Sawmill River (17·0° C) than in the other rivers (15·5, 14·9° C). By influencing community structure and growth of remaining fish, it appears that a strong environmental disturbance can also alter the direction and timing of life histories in Atlantic salmon.     

Early-season river entry of adult Atlantic salmon: its dependency on environmental factors

River entry of adult Atlantic salmon Salmo salar into the River Tornionjoki, monitored during three migration seasons (1997–1999) by horizontal split-beam hydroacoustics, started early in June when water temperature was c . 9° C and when the discharge varied between 1700 and 2000 m³ s⁻¹. In 1997 and 1999, migration peaked during the latter half of June, 17 days after the peak flood, at water temperatures ranging from 11· 5 to 18·2° C. Few statistically significant correlations were observed between river entry and six measured environmental factors and those that were significant were not persistent over the years. The strongest correlation (  r  = −0·60) was between the number of upstream migrants and seawater level, with a time lag of 1 day in 1998. In 1998 and 1999, no clear diurnal migration pattern was observed, although in 1997 the intensity of midday migration was higher than that of the midnight migration. It is concluded that environmental factors have little effect on river entry of Atlantic salmon in a large pristine river located at high latitude.     

Migration of anadromous brown trout Salmo trutta in a Norwegian river

1. Upstream and downstream migrating anadromous brown trout Salmo trutta were monitored daily in fish traps in the River Imsa in south-western Norway for 24 years, from 1976 to 1999. One-third of the fish descended to sea during spring (February–June) and two-thirds during autumn (September–January).
2. In spring, high water temperature appeared to influence the downstream descent. Large brown trout (> 30 cm, chiefly two or more sea sojourns) descended earlier and appeared less dependent on high water temperature than smaller and younger fish. The spring water flow was generally low and of little importance for the descent.
3. In autumn, the daily number of descending brown trout correlated positively with flow and negatively with water temperature.
4. Brown trout ascended from the sea between April and December, but more than 70% ascended between August and October. The number of ascending trout increased significantly with both decreasing temperature and flow during the autumn. This response to flow appeared to be the result of the autumn discharge which is generally high and most fish ascended at an intermediate flow of 7.5–10 m³ s⁻¹ (which is low for the season).
5. In a river like the Imsa with low spring and high autumn flows, water temperature appears to be the main environmental factor influencing the timing and rate of spring descent, while both water temperature and flow seemed to influence the timing and rate of the autumn descent and ascent. These relationships make sea trout migrations susceptible to variation in climate and human impacts of the flow regime in rivers.     

Growth hormone-induced effects on mortality, energy status and growth: a field study on Brown Trout (Salmo trutta)

1. Growth hormone (GH) treatment increases the growth rate and competitive ability of salmonids under laboratory conditions. Since fast growth should increase fitness, why is endogenous secretion of GH not higher in wild fish? To address this question, three hypotheses were suggested. H₁: high GH levels reduce antipredator responses and may therefore increase mortality from predation. H₂: high GH levels reduce long-term (e.g. over winter) survival by reducing allocation to critical energy reserves. H₃: GH is not beneficial for growth under natural conditions.
2. To test these hypotheses, the performance of GH-treated juvenile Brown Trout ( Salmo trutta ) and control (placebo) trout was compared in an enclosed stream section subjected to natural predation. Four experiments were conducted during winter, spring, summer and autumn, respectively.
3. Mortality rates were not significantly different between GH-treated and control trout in any of the four experiments so H₁ was not supported. Energy reserves were generally lower in GH-treated fish, which is consistent with H₂, whereas growth rates in mass were higher in GH-treated fish than in controls so H₃ was not supported. This suggests that GH promotes growth at the expense of investment in maintenance.
4. Judging from growth and mortality rates, the fitness of GH-treated and control trout appeared similar. Thus, escaped GH-manipulated fish may compete successfully with wild fish.
5. Hatchery-raised trout with higher initial condition index suffered higher mortality rates than more slender fish. This novel finding may be explained by reduced escape ability related to body morphology, reduced behavioural responses towards predators by high-condition trout, or predator preferences for high-condition fish.     

Seasonal occurrence of the infectious stage of proliferative kidney disease (PKD) and resistance of rainbow trout, Salmo gairdneri Richardson, to reinfection

The seasonality of proliferative kidney disease (PKD) in rainbow trout, Salmo gairdneri Richardson, at the American River Hatchery, California was found to be primarily dependent on the presence or absence of the infectious stage in the water supply. This was determined by introducing sentinel trout into the hatchery water supply on a monthly basis, followed by their transfer to the laboratory for subsequent holding in 18° C, pathogen-free water for one month prior to examination. These exposures demonstrated that infections were obtained from April through October at ambient temperatures 12–20° C. Trout which had recovered from clinical infections were found to be resistant to reinfection. Resistance was induced by active infection and not just previous exposure to the infectious stage. Trout surviving PKD were also found to harbour later sporogonic stages of the parasite for at least one year following initial infection, but fully formed spores, as judged by well-developed valves, were not observed.     

Factors affecting river entry of adult Atlantic salmon in a small river

In this study, effects of stock origin, fish size, water flow and temperature on time of river ascent of adult Atlantic salmon Salmo salar were tested. Brood stocks were collected in eight Norwegian rivers situated between 59 and 69° N. The fish were reared to smolts, individually tagged and released in the River Imsa, south-west Norway (59° N). Adults from all stocks approached the Norwegian coast concurrently, but Atlantic salmon ≥70 cm in natural tip length entered coastal water slightly earlier during summer than smaller fish. Atlantic salmon <70 cm, however, ascended the river significantly earlier and at lower water flow and higher water temperature than larger fish. Although largest in size, the fish from the northern populations (62–69° N) ascended the River Imsa almost 1 month earlier than those from the south (59–60° N). They seemed less restricted by the environmental factors than the fish originating from the more southern rivers. There was no apparent trend among years in time of river ascent. Maximum ascent per day occurred at water discharges between 12·5 and 15 m³ s⁻¹ and at water temperatures between 10 and 12·5° C. There was a significant positive correlation between water flow and river ascent during the first part of the upstream run from July to September with best correlation for September, when multiple regression analysis indicated that water temperature had an additional positive effect. Stock origin, fish size and water discharge were important variables influencing the upstream migration of Atlantic salmon in small rivers.     

Pools as refugia for brown trout during two summer droughts: trout responses to thermal and oxygen stress

In non–drought years (1977, 1985), temperatures and oxygen concentrations from 1 to 14 July at the deepest point in each of five pools in Wilfin Beck were similar with ranges of 12–18° C and 7·8–9·8 mg l^–1. Trout Salmo trutta were present in all pools. In drought years (1976, 1983), temperature increased and oxygen concentration decreased as pool size decreased. In the two smallest pools, they were outside the thermal and oxygen limits for trout (ranges for both pools 24–29° C, 1·2–2·5 mg l^–1), and trout were absent. Values in a medium–sized pool were close to the incipient lethal levels and a few juvenile trout were present in both drought years. The lowest temperatures and highest oxygen concentrations were recorded in the two largest pools (ranges 20–25° C, 3·6–4·8 mg l^–1) and trout of all ages (0+ to adults) were present in both drought years. In these two pools, both temperature and oxygen concentration decreased from the surface to the deepest point in the pool. Trout preferred lower temperatures near the pool bottom rather than higher oxygen concentrations near the surface, but some fish moved towards the surface at night when the pool cooled slightly. These field results were discussed in relation to lethal values recorded for brown trout in the laboratory, and there was general agreement between field and laboratory values. Trout in the drought years occurred at temperatures close to, or below, the incipient lethal value of 24·7° C (+0·5) and also at the highest oxygen concentrations, but only when these were at temperatures below the incipient lethal value.     

Temperature requirements of Atlantic salmon Salmo salar, brown trout Salmo trutta and Arctic charr Salvelinus alpinus: predicting the effects of climate change

Atlantic salmon Salmo salar, brown trout Salmo trutta (including the anadromous form, sea trout) and Arctic charr Salvelinus alpinus (including anadromous fish) provide important commercial and sports fisheries in Western Europe. As water temperature increases as a result of climate change, quantitative information on the thermal requirements of these three species is essential so that potential problems can be anticipated by those responsible for the conservation and sustainable management of the fisheries and the maintenance of biodiversity in freshwater ecosystems. Part I compares the temperature limits for survival, feeding and growth. Salmo salar has the highest temperature tolerance, followed by S. trutta and finally S. alpinus. For all three species, the temperature tolerance for alevins is slightly lower than that for parr and smolts, and the eggs have the lowest tolerance; this being the most vulnerable life stage to any temperature increase, especially for eggs of S. alpinus in shallow water. There was little evidence to support local thermal adaptation, except in very cold rivers (mean annual temperature <6·5° C). Part II illustrates the importance of developing predictive models, using data from a long-term study (1967-2000) of a juvenile anadromous S. trutta population. Individual-based models predicted the emergence period for the fry. Mean values over 34 years revealed a large variation in the timing of emergence with c. 2 months between extreme values. The emergence time correlated significantly with the North Atlantic Oscillation Index, indicating that interannual variations in emergence were linked to more general changes in climate. Mean stream temperatures increased significantly in winter and spring at a rate of 0·37° C per decade, but not in summer and autumn, and led to an increase in the mean mass of pre-smolts. A growth model for S. trutta was validated by growth data from the long-term study and predicted growth under possible future conditions. Small increases (<2·5° C) in winter and spring would be beneficial for growth with 1 year-old smolts being more common. Water temperatures would have to increase by c. 4° C in winter and spring, and 3° C in summer and autumn before they had a marked negative effect on trout growth.     

Do Norwegian Atlantic salmon feed in the northern Barents Sea? Tag recoveries from 70 to 78° N

Three tagged Atlantic salmon Salmo salar were recaptured as subadults or adults (1·4–3 kg) between 70·5 and 78° N in the western Barents Sea, two of which originated from the Alta Fjord region in northern Norway and one from the Drammen River, south-eastern Norway. An additional tag was recovered from the stomach of a Greenland halibut Reinhardtius hippoglossoides captured south-west of Bear Island at >600 m depth; this tag was from a smolt released in the River Alta 1 month earlier. These are the northernmost tag recoveries reported for Atlantic salmon, and indicate that Norwegian Atlantic salmon, especially the fish from northern populations, may use the northern Barents Sea as a feeding area during part of their life cycle.     

An electrophysiological study on the effects of pH on olfaction in mature male Atlantic salmon (Salmo salar) parr

The olfactory responses of mature male Atlantic salmon ( Salmo salar L.) parr to known odorants were studied after exposure of the olfactory epithelia to water of varying pH. Electrophysiological recordings from the olfactory epithelia indicated that the responses of fish to both testosterone and urine from ovulated female Atlantic salmon were significantly reduced at pH 5·5 and 4·5 and abolished at pH 3·5. Concentration response studies indicated that at pH 5·5 and 6·5 significantly higher concentrations of testosterone and urine were required to produce the same amplitude responses as controls. Both testosterone and urine have previously been shown to be important chemical cues eliciting behavioural and physiological responses in Atlantic salmon. The results are therefore discussed in relation to the possible sublethal effects of acidification on reproduction and behaviour of Atlantic salmon and the effects on salmonid stocks.     

Why,when and how do fish populations decline,collapse and recover? The example of brown trout (Salmo trutta) in Rio Chaballos (northwestern Spain)

1. Around the year 2000, historically abundant populations of brown trout ( Salmo trutta ) and Atlantic salmon ( S. salar ) co-occurring in rivers flowing along the Cantabrian corridor of north-western Spain showed a dramatic decline to alarming levels. For one reason or another, fishing was not banned and fishing pressure continued to reduce the few survivors. Unexpectedly, the populations recovered 'naturally' in a very short time period to the extent that in 2004–06 numbers had attained population sizes comparable to those of the mid 1980s.
2. The population of brown trout in Rio Chaballos showed a boom-and-bust pattern concurrent with those observed across broader geographical scales. This study revisits a 22-year data set to explore the nature of the severe decline and rapid recovery of this population.
3. Recruitment was related to stream discharge in March that covers the emergence period and the earliest search stages for food. Coefficients of variation for discharge and recruitment increased over the years and were highly correlated with each other, demonstrating that increased temporal variability in recruitment is strongly linked to increased variability in stream discharge. In turn, recruitment appears to be the major determinant of year-class strength and hence, of population size.
4. A number of factors appeared to operate as resilience mechanisms, permitting the population to increase rapidly when environmental conditions are optimal. These include strong recruitment-discharge relationships, short life-span, small stocks of eggs or offspring required to fill the amount of space suitable for the youngest juveniles and a few females surviving to spawn for a second time.
5. Implications for fishery management and conservation are discussed in the context of the expectation that these populations will vary over the coming years within ranges similar to those in this study.     

Daylight responses to overhead cover in stream channels for fry of four salmonid species

Brown trout ( Salmo trutta ), Atlantic salmon ( Salmo salar ), brook trout ( Salvelinus fontinalis ) and lake trout ( Salvelinus namaycush ) fry entering the free-feeding stage, were tested for overhead cover preferences in stream channels at different temperatures and water velocities. Atlantic salmon showed strong preferences for overhead cover, brown trout moderate preferences, whereas lake trout had preferences only at high temperatures, i.e. 12.4–19.2°C. Brook trout showed no cover preferences. Temperature influenced cover preferences of Atlantic salmon and brown trout considerably. The fry tended to seek more cover at low temperatures, i.e. 6.0–8.3°C     

Survival and growth of brown trout Salmo trutta L. embryos and the timing of hatching and emergence in two boreal lake outlet streams

Survival, growth and hatching of brown trout Salmo trutta embryos were studied using in situ incubation experiments in two lake outlet streams in Finland. The experimental design in both streams included an outlet site and a reference site far downstream. The date of hatching was recorded and the Elliott–Hurley model was then used to predict the time of emergence based on water temperature. For data analyses, the incubation period was divided into 'winter' (from fertilization to mid March) and 'spring' (from mid March until the end of the experiment). Temperature of the large-lake outlet remained at 1° C through the winter, while in other sites temperature was close to 0° C. In spring, temperature increased more slowly in the large-lake outlet. The survival of embryos was overall very high, from 83 to 98%, and they were larger in the outlets than in the downstream sites. Embryos hatched at the large-lake outlet in March, and 3–5 weeks later in the other sites. Although there were considerable between-site differences in hatching intervals, difference in expected 50% emergence dates of the earliest and latest site was only 4 days. Thus, any growth advantage that the outlet embryos had in winter disappeared by the end of the alevin period. Lake outlets, however, may be important for age 0 year brown trout later during the summer when other stream habitats do not provide adequate food resources.     

Mortality in Atlantic salmon (Salmo salar) associated with trichodinid ciliates

A protozoan infection (Trichodina truttae) was identified in captive Atlantic salmon (Salmo salar) kelts that died in spring of 1988 and 1989. Fish with intense infections showed signs of listlessness, erratic swimming and inappetence. The infection induced excessive mucus secretion, epithelial sloughing and lesions that probably permitted entry of opportunistic bacteria which eventually caused ulcers and death. A seawater bath for 30 min each week for 4 wk effectively controlled the parasite.     

Sea growth, smolt age and age at sexual maturation in Atlantic salmon

Relationships between growth at sea, smolt size and age at sexual maturation of Atlantic salmon Salmo salar were tested. The fish were offspring of brood stocks sampled in eight Norwegian rivers at latitudes between 59° and 70° N, hatchery reared and released at smolting at the mouth of the River Imsa (59° N). Smolt size influenced the subsequent growth rate of Atlantic salmon. The larger the fish were at release, the slower the yearly length increment at sea. Mean sea age at sexual maturity, measured as proportion of the returning adults attaining sexual maturity at sea age 2 years, was significantly correlated with mean growth rate during the first year at sea and mean smolt size ( r ²= 0·74, P < 0·001). Fish attaining maturity at a relatively high sea age were more fast growing during their first year at sea than those maturing at a younger age. The results indicate that high sea age at sexual maturation is a population-specific characteristic and associated with high early growth rate at sea.     

Body temperature and seawater adaptation in farmed Atlantic salmon and rainbow trout during prolonged chilling

The core temperature of the rainbow trout Oncorhynchus mykiss (3·5 kg) dropped to 1·0° C during the first 6 h of chilling at 0·5° C, remained stable until 24 h, and dropped significantly to 0·7° C after 39 h. Blood plasma osmolality increased and muscle moisture content decreased gradually with increasing chilling time. After 39 h of chilling, the rainbow trout experienced 40 mosmol l^-1 higher blood plasma osmolality and 2·8% less muscle moisture content compared with initial values. In the Atlantic salmon Salmo salar (5·3 kg), core temperature dropped to 1·3° C and blood plasma osmolality increased significantly during the first 6 h of chilling at 0·5° C, but remained relatively stable throughout the rest of the experimental period. After 39 h of chilling, the salmon experienced 20 mosmol l^-1 higher blood plasma osmolality and 0·5% less muscle moisture content compared with initial values. In rainbow trout muscle moisture content was inversely related to blood plasma osmolality indicating reduced seawater adaptation with increasing hours of chilling. No such relationship was observed in the Atlantic salmon. Hence, changes in plasma osmolality and muscle moisture in the Atlantic salmon do not indicate osmoregulatory failure since the new levels, once established, were maintained throughout the chilling time.     

Age, sex ratio and timing of the catch of kelts and ascending Atlantic salmon in the subarctic River Teno

By 15 June, 82% of the catch of Atlantic salmon Salmo salar kelts had been taken from the middle part of River Teno, northern Scandinavia. The median date of capture was 4 June for males and 8 June for females. Salmon of 1–4 sea–winters (SW) of both sexes survived spawning to return to sea as kelts. Among males, 1 SW kelts were caught earliest in the spring and 3 SW latest, but among females 4 SW were earliest, then 3 SW and finally 1 and 2 SW. There were 17 river and sea–age combinations among the kelts compared with 23 among the ascending salmon. The smolt age distribution and the mean smolt age differed significantly only between female 2 SW ascending salmon (3·97 years) and kelts (4·14 years). The proportion of 1 SW females was higher and that of 3 SW males lower among kelts than among ascending salmon. The proportion of males among 1 SW ascending salmon was 80% but among kelts only 57%. Similarly, the proportion of males among 3 SW fish was 21% for ascending salmon but only 7% for kelts. Hence overwinter mortality was higher among males. Male and female kelts of 1 and female kelts of 2 SWhad a greater mean length than ascending salmon in corresponding groups indicating a better survival of larger fish within an age group. Grilse ascend rivers after most kelts have left, but the main catch of ascending 2–3 SW salmon takes place concurrently with kelts leaving the river, inadvertently targeting kelts in the fishery.     

Adaptive and Neutral Genetic Variation and Colonization History of Atlantic Salmon, Salmo salar

Synopsis I combined neutral microsatellite markers with the major histocompatibility complex (MHC) class IIB to study genetic differentiation and colonization history in Atlantic salmon, Salmo salar, in the Baltic Sea and in the north-eastern Atlantic. Baltic salmon populations have lower levels of microsatellite genetic variation, in terms of heterozygosity and allelic richness than Atlantic populations, confirming earlier findings with other genetic markers, suggesting that the Baltic Sea populations have been exposed to genetic bottlenecks, most likely at a founding event. On the other hand, the level of MHC variation was similar in the Baltic and in the north-eastern Atlantic, indicating that positive balancing selection has increased the level of MHC-variation. Both microsatellite and MHC class IIB genetic variation give strong support to the hypothesis that the Baltic salmon are of a biphyletic origin, the southern population in this study is strongly differentiated from both the northern Baltic salmon populations and from the north-eastern Atlantic populations. Salmon may have colonized the northern Baltic Sea either from the south, via the so called “N?rke strait” or from the north, via a proposed historical connection between the White Sea and the northern Baltic. At microsatellites, no significant isolation-by distance was found at either colonization route. At the MHC, populations were significantly isolated by distance when assuming that colonization occurred via the “N?rke strait”.     

Survival data for dewatered Rainbow Trout (Salmo gairdneri Rich.) eggs and alevins

Survival of four intergravel development phases of rainbow trout, Salmo gairdneri Rich., in a dewatered environment varied with stage of development, duration of exposure, and relative humidity. Egg phases were considerably more tolerant than alevin phases. Cleavage eggs tolerated dewatering for 12 consecutive days (97% survival), the maximum period tested. Embryos tolerated dewatering until they began hatching on the eight day, when mortalities sharply increased. In contrast, eleutheroembryos had only 43% survival when dewatered 6h and pre-emergent alevins had only 36% survival when dewatered 2h. A few dewatered embryos died when relative humidity was 100%, but mortality increased significantly at 90 and 80% relative humidity. Survival of alevin phases, but not egg phases, in residual, non-flowing water was limited by formation of oxygen deficits.