Short‐term freshwater exposure benefits sea lice‐infected Atlantic salmon

Atlantic salmon Salmo salar were infected with sea lice Lepeophtheirus salmonis (0·08 ± 0·007 sea lice g⁻¹) over a period of 4 h. Both infected and non‐infected fish were swim tested in sea water (SW) and fresh water (FW). The ventral aorta of each fish was fitted with a Doppler cuff in order to measure cardiac output, stroke volume and heart rate during swim testing. Blood samples were taken at rest and after exercise. Critical swimming speed of infected fish in SW (2·14 ± 0·08 body lengths, bl s⁻¹) was significantly lower ( P  < 0·05) than infected fish switched to FW (2·81 ± 0·08 bl s⁻¹) and non‐infected fish in SW (2·42 ± 0·04 bl s⁻¹) and FW (2·61 ± 0·08 bl s⁻¹). Cardiac and blood results indicated infected fish exposed to FW did experience stress, but relief from osmotic and ionic distress probably reduces energy expenditure, allowing the increase in performance. As the performance of sea lice‐infected fish improved upon transfer to FW, it is likely that heavily infected salmonids do return to FW to restore compromised osmotic and ionic balance, and remove sea lice in the process.     

Return migration of one‐sea‐winter Atlantic salmon in the River Tana

Fifty‐three one‐sea‐winter Atlantic salmon Salmo salar (45–63 cm L _T) were radio‐tagged in the Tana fjord, Barents Sea, in 1995. Thirty‐seven fish (70%) entered the freshwater zone of the River Tana in an average of 3 days after release in the fjord. The migration speeds in the lowest river section below the first riffle area were significantly higher than in the subsequent river section below the second riffle area. Similarly, the observed time spent in the first riffle area was significantly lower than in the next riffle area. The majority of Atlantic salmon entered the river during the hours of high tide and the subsequent ebb tide. In addition, most river entries were recorded around midnight. No effects of river flow on the river entry or migration speed were detected, but the migration speed of Atlantic salmon in both river sections examined was greater at lower temperatures. Twenty‐eight fish (72%) were recaptured in the river, 71% of them with weirs and gillnets, and 29% by rod and line. Over half of the Atlantic salmon (54%) were recaptured within 3 weeks following river entry, and within the first 100 km of the river (56%). The results are discussed in relation to earlier studies on multi‐sea‐winter Atlantic salmon in the River Tana.     

Physiological impact of sea lice on swimming performance of Atlantic salmon

Atlantic salmon Salmo salar were infected with two levels of sea lice Lepeophtheirus salmonis (0·13 ± 0·02 and 0·02 ± 0·00 sea lice g⁻¹). Once sea lice became adults, the ventral aorta of each fish was fitted with a Doppler cuff to measure cardiac output ( ̇ ), heart rate ( f _H) and stroke volume ( V _S) during swimming. Critical swimming speeds ( U _crit) of fish with higher sea lice numbers [2·1 ± 0·1 BL (body lengths) s⁻¹] were significantly lower ( P  < 0·05) than fish with lower numbers (2·4 ± 0·1 BL s⁻¹) and controls (sham infected, 2·6 ± 0·1 BL s⁻¹). After swimming, chloride levels in fish with higher sea lice numbers (184·4 ± 11·3 mmol l⁻¹) increased significantly (54%) from levels at rest and were significantly higher than fish with fewer lice (142·0 ± 3·7 mmol l⁻¹) or control fish (159·5 ± 3·5 mmol l⁻¹). The f _H of fish with more lice was 9% slower than the other two groups at U _crit. This decrease resulted in ̇ not increasing from resting levels. Sublethal infection by sea lice compromised the overall fitness of Atlantic salmon. The level of sea lice infection used in the present study was lower than has previously been reported to be detrimental to wild Atlantic salmon.     

Endurance swimming of diploid and triploid Atlantic salmon

When groups of diploid (mean ±  s . e . fork length, L _F) 33·0 ± 1·4 cm and triploid (35·3 ± 0·5 cm) Atlantic salmon Salmo salar were forced to swim at controlled speeds in a carefully monitored 10 m diameter 'annular' tank no significant difference was found between the maximum sustained swimming speeds ( U _ms, maintainable for 200 min) where the fish swam at the limit of their aerobic capability. Diploids achieved 2·99 body lengths per second (bl s⁻¹)(0·96 m s⁻¹) and triploids sustained 2·91 bl s⁻¹(1·02 m s⁻¹). The selection of fish for the trials was based on their ability to swim with a moving pattern projected from a gantry rotating at the radius of the tank and the selection procedure did not prove to be significant by ploidy. A significant difference was found between the anaerobic capabilities of the fish measured as endurance times at their prolonged swimming speeds. During the course of the experimentation the voluntary swimming speed selected by the fish increased and the schooling behaviour improved. The effect of the curvature of the tank on the fish speeds was calculated (removing the curved effect of the tank increased the speed in either ploidy by 5·5%). Implications of the endurance times and speeds are discussed with reference to the aquaculture of triploid Atlantic salmon.     

Evaluation of biochemical methods for the non‐destructive identification of sex in upstream migrating salmon and sea trout

Female‐specific markers of reproductive activity [plasma 17β‐oestradiol (E2), vitellogenin (VTG) and alkali‐labile phosphoprotein phosphorous (ALP)] were measured over 12 months in a captive population of brown trout Salmo trutta . During the early months of the reproductive season (February to May) and using the concentration of plasma E2 or plasma ALP as a marker for females the proportion of fish in which sex was misidentified was high (15–50%). The misidentification rate was considerably lower (1–8%) using plasma VTG. Preliminary evaluation of a commercial immunochromatographic VTG test system as a screen for the presence or absence of VTG in plasma from brown trout provided results that were consistent with those obtained from direct measurement of plasma VTG levels by enzyme‐linked immunosorbent assay (ELISA). These preliminary conclusions were verified by sampling upstream‐migrating anadromous brown trout, sea trout, and Atlantic salmon Salmo salar trapped over a 6 month period. Plasma E2 levels did not satisfactorily discriminate between male and female sea trout and Atlantic salmon. Plasma VTG levels in both species, however, were bimodally distributed and it was assumed that this divergence corresponded to male (plasma VTG levels <10 μg ml⁻¹) and female (plasma VTG levels >800 μg ml⁻¹) fishes. Plasma ALP provided a more accurate indication of sex in the wild Atlantic salmon and sea trout than was suggested by the pilot study on captive brown trout. The commercial immunochromatographic VTG test system provided results that were wholly consistent with the data obtained from the trapped fishes by direct measurement of plasma VTG.     

Comparing upriver spawning migration of Atlantic salmon Salmo salar and sea trout Salmo trutta

Radio tagged wild Atlantic salmon Salmo salar(n = 30) and sea trout Salmo trutta(n = 19) were simultaneously released from a sea pen outside the mouth of the River Lærdalselva and their migration to spawning areas was recorded. The distance from the river mouth to a position held at spawning ranged from 2 to 24 km and did not differ between the species (mean ± s .d . 15·9 ± 4·3 and 14·9 ± 5·2 km for Atlantic salmon and sea trout, respectively). The duration of the migration phase, however, was significantly shorter for Atlantic salmon than for sea trout (8–12 days, respectively). All Atlantic salmon migrated straight to an area near the spawning ground, whereas 50% of the sea trout had a stepwise progression with one or more periods with erratic movements before reaching the spawning area. After the migration phase, a distinct search phase with repeated movements up‐ and downstream at or close to the position held at spawning was identified for the majority of the fishes (75%, both species). This search phase was significantly shorter for Atlantic salmon than for sea trout (mean 13–31 days, respectively). Mean ± s .d . length of the river stretch used during the search phase was larger for sea trout (3·3 ± 2·5 km) than for Atlantic salmon (1·2 ± 0·9 km). A distinct holding phase, with no movements until spawning, was also observed in the majority of the Atlantic salmon (80%, mean duration 22 days) and sea trout (65%, mean duration 12 days). For both species, a weak, non‐significant trend was observed in the relationship between time spent on the migration phase, and time spent on the search (r² = 0·43) and holding phase (r² = 0·24). There was a highly significant decrease, however, in the duration of the holding phase with an increase in the time spent on the search phase (r² = 0·67).     

Reproductive migration of brown trout in a small Norwegian river studied by telemetry

The movement of 34 large (39–73 cm standard length) brown trout Salmo trutta was monitored using radio telemetry for up to 74 days in Brumunda, a small Norwegian river (mean annual discharge 3·3 m³ s⁻¹) flowing into the large Lake Mjøsa. The maximum range of movement in the river was 20 km. No clear relationships existed between individual movement and water discharge, temperature and barometric pressure. Brown trout migrated at all levels of water discharge. At low discharge (<2 m³ s⁻¹) movements were nocturnal. A weir 5·3 km from the outlet restricted ascending brown trout at low ( c . 6° C), but not at high ( c . 8° C) water temperatures. Spawning occurred in September to October and tagged individuals spent 2–51 days at the spawning sites. Mean migration speed from tagging to when the fish reached the spawning area, and from when they left the spawning areas and reached the lake was 1·0 and 2·3 km day⁻¹, respectively. All tagged brown trout that survived spawning returned to the lake after spawning.     

Predator‐induced hyperventilation in wild and hatchery Atlantic salmon fry

Following exposure to a predator stimulus (a brown trout Salmo trutta ), the opercular rate of Atlantic salmon Salmo salar fry increased by 35·3 ± 11·0%(mean ± 95% CI). The time taken for opercular rate to decline to baseline levels depended upon the occurrence of any associated locomotory activity. Opercular rates of fish that dashed when exposed remained elevated for 38·2 ± 20·6 min, whereas those of individuals that did not move ('freezers') recovered within 7·2 ± 2·9 min. The duration that opercular rate remained elevated was positively correlated with the magnitude of the elevation, which was higher in 'dashers' than freezers. The maximum opercular rate in 'freezers' was similar between wild fry and hatchery‐reared fry (from wild parents). There was a significant delay, however, in hatchery compared with wild fry in the time until peak ventilatory response and onset in the decline phase. This difference in opercular response suggests that hatchery fish were slower to realize fully the potential danger from the predator. Any delay in response could be directly attributed to the effect of hatchery‐rearing environment, rather than domestication or hatchery selection effects.     

Dissolved oxygen concentration and emergence of sea trout fry from natural redds in tributaries of the River Rhine

A total of 4289 sea trout Salmo trutta alevins swam up from six of eleven covered redds. They emerged only from those redds where the mean dissolved oxygen level was >6·9 mg l⁻¹ and the Sorting coe.cient, an indicator of intragravel void space, was <5. These results are discussed in relation to field studies using artificially buried salmonid eggs but also in relation to the ongoing programme for the reintroduction of Atlantic salmon in tributaries of the River Rhine.     

Effect of increased flow on the behaviour of Atlantic salmon parr in winter

The effect of increased flow on movement and microhabitat use of Atlantic salmon Salmo salar parr in winter was investigated using radiotelemetry. To simulate hydropeaking operations, flow was increased four‐fold from 1·3 m³ s⁻¹ to 5·2 m³ s⁻¹ for 24 h periods. Flow did not affect fish habitat use or displacement and had little effect on fish activity within diel periods. During high flow periods in late winter, fish reduced night‐time activity. Stranding rates during flow reduction were also very low (only one fish).     

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 speeds and orientation of Atlantic salmon and sea trout post-smolts in a Norwegian fjord system

We recorded the observed and actual swimming speeds of Atlantic salmon and sea trout post-smolts in a Norwegian fjord system, and initiated studies on the orientation mechanisms of the post-smolts. We tracked Atlantic salmon and sea trout with acoustic transmitters for up to 14 h after release. The actual swimming speed and direction of a fish relative to the ground is the vector sum of the observed movements of the fish and the movements of the water. We determined actual swimming speeds and directions of the post-smolts, which reflect their real swimming capacities and orientation, by corrections for the speed and direction of the water current. The post-smolts were actively swimming. The observed direction of movement was dependent on the actual movement of the fish and not the water current. Water currents were not systematically used as an orientation cue either in Atlantic salmon or sea trout, as the actual movements were random compared to the direction of the water current. The actual movement of sea trout were in all compass directions, with no systematic pattern. The Atlantic salmon also moved in all compass directions, but with the lowest frequency of actual movement towards the fjord.     

Fjord migration and survival of wild and hatchery-reared Atlantic salmon and wild brown trout post-smolts

The behaviour of wild (n = 43, mean L_T = 152 mm) and hatchery-reared (n = 71, mean L_T = 198 mm) Atlantic salmon and wild anadromous brown trout (n = 34, mean L_T = 171 mm) post-smolts with acoustic transmitters was compared in a Norwegian fjord system. There was no difference in survival between wild and hatchery reared salmon from release in the river mouth to passing receiver sites 9.5 km and 37.0 km from the release site. Mortality approached 65% during the first 37 km of the marine migration for both groups. There was no difference between wild and hatchery-reared salmon either in time from release to first recording at 9.5 km (mean 135 and 80 h), or in the rate of movement through the fjord (mean 0.53 and 0.56 bl s⁻¹). Hatchery-reared salmon reached the 37 km site sooner after release than the wild salmon (mean 168 and 450 h), but rate of movement in terms of body lengths per second did not differ (mean 0.56 and 0.77 bl s⁻¹). The brown trout remained a longer period in the inner part of the fjord system, with much slower rates of movement during the first 9.5 km (mean 0.06 bl s⁻¹).     

The survival of semi‐wild, wild and hatchery‐reared Atlantic salmon smolts of the Simojoki River in the Baltic Sea

The recapture rate and survival of hatchery‐reared Atlantic salmon Salmo salar stocked as 1 year‐old parr (semi‐wild) with that of hatchery‐reared Atlantic salmon stocked as 2 year‐old smolts and wild smolts of Atlantic salmon in the northern Baltic Sea were compared. This was done through tagging experiments carried out in 1986–1988 and 1992. The recapture rate of the semi‐wild groups varied from 1·0 to 13·1%, being similar in 3 tagging years and lower in 1 year than that of the wild groups (1·7–17·0%). The recapture rate of the semi‐wild groups was similar (in 2 years) or higher (in 2 years) than that of the hatchery‐reared groups stocked as smolts (1·3–6·3%). The survival of semi‐wild smolts during the sea migration was as high as that of wild Atlantic salmon of an equal size and two to three times higher than hatchery‐reared Atlantic salmon stocked as smolts. The survival rate was positively associated with smolt size. The suitability of hatchery‐reared parr and smolts in the management of reduced Atlantic salmon stocks is compared.     

Effects of salmon lice infection and salmon lice protection on fjord migrating Atlantic salmon and brown trout post-smolts

Effects of artificial salmon lice infection and pharmaceutical salmon lice prophylaxis on survival and rate of progression of Atlantic salmon (n = 72) and brown trout post-smolts (n = 72) during their fjord migration, were studied by telemetry. The infected groups were artificially exposed to infective salmon lice larvae in the laboratory immediately before release in the inner part of the fjord to simulate a naturally high infection pressure. Groups of infected Atlantic salmon (n = 20) and brown trout (n = 12) were also retained in the hatchery to control the infection intensity and lice development during the study period. Neither salmon lice infection nor pharmaceutical prophylaxis had any effects on survival and rate of progression of fjord migrating Atlantic salmon post-smolts compared to control fish. Atlantic salmon spent on average only 151.2 h (maximum 207.3 h) in passing the 80 km fjord system and had, thus, entered the ocean when the more pathogenic pre-adult and adult lice stages developed. The brown trout, in comparison to Atlantic salmon, remained to a larger extent than Atlantic salmon in the inner part of the fjord system. No effect of salmon lice infection, or protection, was found in brown trout during the first weeks of their fjord migration. Brown trout will, to a larger extent than Atlantic salmon, stay in the fjord areas when salmon lice infections reach the more pathogenic pre-adult and adult stages. In contrast to Atlantic salmon, they will thereby possess the practical capability of returning to freshwater when encountering severe salmon lice attacks.     

Is Atlantic salmon production limited by number of territories?

Thirty Atlantic salmon Salmo salar parr (mean 135 mm total length, L _T) in the River Alta, Norway, were radio‐tagged and tracked during a 11–13 day period. The parr stayed within defined home ranges with a 95% probability of localization within an average area of 1286 m²(241–3484 m²). On average, each parr had overlapping home ranges with 7·7 other parr, and the overlap between pairs of fish covered on average 24% of their home range areas. Mean length of the river stretch used was 90 m (22–383 m). On average, during 38%(16–77%) of the tracking surveys, the fish had moved >10 m since the previous survey. Mean total distance moved during the whole study was 402 m (208–862 m). The Atlantic salmon were most often recorded in riffles, but 27% of the parr alternated between riffles and pools. The extensive movements, flexible habitat utilization and relatively large home ranges, together with the fact that all the parr had home ranges partly overlapping with other radio‐tagged parr, indicate a flexible and variable behaviour and, thereby, a more complex social structure than a rigid defence of a fixed territory. Thus, Atlantic salmon production seems not to be limited by the highest potential number of territories for large parr in a given area.     

Seasonal changes in the body composition of young riverine Atlantic salmon and brown trout

The body composition of protein and fat in Atlantic salmon Salmo salar and brown trout Salmo trutta before and after winter was investigated in a temperate, small river, normally ice covered from the middle of November until the end of March. Fat, protein and specific energy declined greatly in winter but were replenished rapidly in spring. Rates of decline were slower for the smallest fish, which also had the lowest specific content of fat, protein, and energy, while the differences in absolute amounts were greatest for the largest fish. The mean specific fat content was reduced by 45–70% during winter, relative to the pre-winter period (September). Mean daily reductions in specific enegy of the larger size groups of brown trout (3·7 × 10⁻³ kJ g⁻¹ day⁻¹) were almost half of the corresponding values for the largest Atlantic salmon (6·3 × 10⁻³ kJ g⁻¹ day⁻¹) during winter. A minor reduction in protein content was found during winter, with mean reductions of 6–10% in comparison to those in September. During spring the fat content was replenished rapidly, particularly for the smallest salmon fry (a threefold increase from April to June). Fat content in the larger salmon and trout increased by about 1·8 times. Based on estimated metabolic rates, digested energy during wintertime may contribute about two-thirds of the brown trout fry's energy demand. For Atlantic salmon, the corresponding value is about 50%. The winter period put considerable stress on the young salmonids living in lotic environments, in particular for the smallest fry with the lowest energy content before winter and the largest losses during winter. This should make the fry more vulnerable to adverse abiotic and biotic factors.     

Physiological characteristics of wild Atlantic salmon post-smolts during estuarine and coastal migration

Changes were measured in some of the major physiological variables associated with seawater adaptability, growth and energetics in wild Atlantic salmon Salmo salar smolts and post-smolts migrating from the river and through the estuary, fjord and coastal areas in the River Orkla and the Trondheimsfjord, Norway during late May to early June. Gill Na⁺,K⁺-ATPase activity increased to levels of 12–16 µmol ADP mg protein⁻¹ h⁻¹ in post-smolts caught in higher salinity zones, probably representing long-term levels of Atlantic salmon post-smolts in oceanic conditions. Muscle moisture was regulated within narrow limits (77·7–78·7%) in fish from all zones during both years, suggesting that post-smolts adapt to marine conditions without any long-term disturbance of hydro-mineral balance. Lipid and glycogen content showed a general trend towards depletion from the river, through the fjord and into the ocean. There was, however, no significant change in protein content. The present results confirm that smolts are naturally 'energy deficient' during downstream migration, and suggest that post-smolts also mobilize energy reserves during their early marine phase, while protein is allocated for somatic growth. Plasma growth hormone (GH) levels increased transiently during passage through the estuary and fjord, with lower levels observed in post-smolts caught off-shore, i.e. in fish which were feeding on marine prey and had adapted to the marine environment. These physiological changes may confer substantial selective advantages during the critical early marine phase of anadromous salmonids, and hence are adaptive for long-term survival in sea water.     

Stream channel experiments on downstream movement of recently emerged trout, Salmo trutta L. and salmon, S. salar L.—I. Effect of four different water velocity treatments upon dispersal rate

Four experimental stream channels were used to study instantaneous downstream dispersal rates of young trout, Salmo trutta L., and salmon, S. salur L ., relative to four different water velocities.
Young salmon showed a high rate of dispersal at a low velocity of 7.5 cm s⁻¹ and lower rates at higher velocities of 25 to 70cm s⁻¹. Trout showed their lowest rate at 25cm s⁻¹ with a slightly higher rate at 7.5 cm s⁻¹ and increasingly higher rates at velocities in excess of 25 cm s⁻¹. These results are consistent with field observations on the velocity preferences of young trout and salmon.     

Seasonal occurrence of sea lice Lepeophtheirus salmonis on sea trout in two north Norwegian fjords

Seasonal occurrence of the parasitic copepod Lepeophtheirus salmonis (sea lice) was studied from March to December 2001 in two large north Norwegian sill fjords without fish farming activity, the Ranafjord and the Balsfjord. Anadromous brown trout Salmo trutta (sea trout) in both fjords had a low infestation rate during all sampling periods, but followed a seasonal pattern. During early and late winter (November to December and March to April) and spring (May to June), the prevalence varied from 0 to 25% and the abundance was <0·5 sea lice. Adults dominated (92%) during this period, particularly gravid females. In both fjords, the highest prevalence was during September (80–81%, all stages represented). In Ranafjord, the abundance and mean intensity during this month was 6·8 and 8·6 sea lice, respectively, while in Balsfjord it was 3·6 and 4·5 sea lice, respectively. Fish were captured at temperatures down to 1° C and at full strength sea water which is supposed to cause osmoregulatory problems for the fish. This observation has implications for the understanding of high‐latitude sea trout behaviour and can also make the fish more vulnerable to heavy sea lice infestation during this period. It is suggested that winter running sea trout help to maintain a self replicating local population of sea lice within such fjord systems where other possible hosts ( e.g . farmed Atlantic salmon Salmo salar ) are not present during a whole year cycle.