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.     

Feeding ecology of early marine phase Atlantic salmon Salmo salar post-smolts

Dietary analyses of Atlantic salmon Salmo salar post-smolt stomachs collected from 2001 to 2005 in Penobscot Bay, Maine, U.S.A., have yielded insights into the feeding ecology of early marine phase post-smolts from different rearing origins. Most stomachs contained only one or two prey types, suggesting active prey selection. Post-smolts that lived in the river longer (i.e. from naturally reared and parr-stocked origins) were smaller and consumed more fishes than invertebrates compared to larger post-smolts that emigrated immediately post-stocking (i.e. from smolt-stocked origins). Naturally reared S. salar consumed c. 84% fishes and 16% crustaceans and parr-stocked S. salar consumed 64% fishes and 34% crustaceans. Stocked smolts consumed 48% fishes and 40% crustaceans. Differences in the type and quantity of consumed prey may be indicative of behavioural differences among rearing origins that influence post-smolt survival.     

Interaction between migration behaviour and estuarine mortality in cultivated Atlantic salmon Salmo salar smolts

Migration behaviour and estuarine mortality of cultivated Atlantic salmon Salmo salar smolts in a 16 km long estuary were studied using two methods: (1) acoustic telemetry and (2) group tagging in combination with trap nets. Progression rates of surviving individuals through the estuary were relatively slow using both methods [0·38 L_T (total length) s^?1 v. 0·25 L_T s^?1]. In 2012, the progression rate was slow from the river to the estuary (0·55 L_T s^?1) and the first part of the estuary (0·31 L_T s^?1), but increased thereafter (1·45–2·21 L_T s^?1). In 2013, the progression rate was fast from the river to the estuary (4·31 L_T s^?1) but was slower thereafter (0·18–0·91 L_T s^?1). Survival to the fjord was higher in 2012 (47%) compared to 2013 (6%). Fast moving individuals were more likely to migrate successfully through the estuary compared to slower moving individuals. Adult recapture of coded‐wire‐tagged S. salar was generally low (0·00–0·04%). Mortality hot spots were related to topographically distinct areas such as the river outlet (in 2012) or the sill separating the estuary and the fjord (in 2013). At the sill, an aggregation of cod Gadus morhua predating on cultivated smolts was identified. The results indicate that slow progression rates through the estuary decreases the likelihood of smolts being detected outside the estuary. The highly stochastic and site‐specific mortality patterns observed in this study highlight the complexity in extrapolating mortality patterns of single release groups to the entire smolt run of wild S. salar.     

Reproductive strategies of Atlantic salmon: ecology and evolution

Atlantic salmon (Salmo solar, Salmonidae) show a diversity of life history, behavioural and morphological adaptations for reproduction which have evolved as an outcome of competition to maximize reproductive success. Reproductive traits of females have been shaped principally by natural selection for offspring production and survival, those of males by sexual selection for access to matings. Female Atlantic salmon invest approximately six times more energy in offspring production (i.e. gonads) than males and face an important trade-off between number and size of eggs to produce that will maximize the number of surviving offspring. Timing of breeding and the construction of nests appear adapted to increase offspring survival. The most important determinant of female breeding success is body size because it affords high fecundity, access to breeding territories and decreased probability of nest destruction. Asynchronous female spawning and the male ability to spawn rapidly and repeatedly results in male-biased operational sex ratios that generate intense male competition for mates. This has likely been responsible for the evolution of elaborate male secondary sexual characters associated with fighting and status signalling. Furthermore, it has given rise, through frequency-dependent selection, to two alternative male breeding phenotypes: (1) large, anadromous males; and (2) small, mature male parr. Anadromous males invest heavily in behavioural activity on the spawning grounds, searching and fighting for mates and courting them, with body size being an important determinant of their breeding success. This behavioural activity carries a heavy cost, as anadromous males have significantly reduced survival relative to females. In contrast, mature male parr invest proportionally more in testes for sperm competition and attempt to sneak access to matings. While this behaviour also carries costs in terms of subsequent growth and survival, male parr are more likely to breed again, either prior to or following a migration to sea, than anadromous males. While knowledge about the breeding of Atlantic salmon is detailed, we are only beginning to understand the ultimate causes and/or functional significances of their reproductive strategies. Predictive models of the life history variation are developing, focusing on the need for empirical study and testing of life history and reproductive patterns.     

Rhythmic aspects of estuarine migration of hatchery-reared Atlantic salmon (Salmo salar) smolts

Seasonal, diel and tidal rhythmic activity of hatchery-reared Atlantic salmon ( Salmo salar ) smolts migrating through a large estuary was studied by ultrasonic tracking of 46 individuals during two seasons. Prior to 10 May each year most smolts were inactive and remained near shore in shallow water. After 10 May nearly all smolts moved away from the release point into swift water and made rapid seaward progress. Initiation of migration each year occurred when river and hatchery water temperatures rose above 9°C. Migration in the estuary was largely passive drift, and as a result there were distinct tidal rhythms of ground ('swimming') speed and net seaward travel. There were no diel rhythms in ground speed or net seaward travel; smolts drifted seaward on the tides during both day and night. Smolts may be slightly deeper during day than night.     

A size-dependent migration strategy in Atlantic salmon smolts: Small smolts favour nocturnal migration

Migration theory states that migration behavioural strategies should be optimised to maximise fitness. Many studies have shown that in downstream migrating Atlantic salmon Salmo salar L. smolts, mortality from predation is high and negatively size dependent. The most common predators are birds and piscivorous fish that are mainly daylight feeders. Given the high mortality during this stage we should expect to observe smolts to follow predator avoidance strategies that may be affected by body size. We tested the hypothesis that small smolts have a higher tendency to exhibit predator avoidance strategies (i.e. nocturnal versus diurnal migration) than larger smolts. The number and size of out-migrating/downstream-migrating wild Atlantic salmon smolts was recorded as they passed through a glass-sided channel during April-May, 1996–1999. In all years, the mean size of nocturnal migrating smolts was significantly lower than the mean size of diurnal migrating smolts. Analysis of the size of smolts, during early and late stages of the migration period showed size-dependent nocturnal migration behaviour up to the end of April. After this, no such size dependent migration pattern was observed. However, small smolts (<100 mm) were absent during this period. We suggest that nocturnal migration is an adaptive behaviour that small Atlantic salmon smolts have to avoid predation by large daylight feeding visual piscivorous predators (e.g. pike Esox Lucius L. and fish eating birds).     

The North Atlantic subpolar gyre and the marine migration of Atlantic salmon Salmo salar: the ‘Merry‐Go‐Round’ hypothesis

One model for marine migration of Atlantic salmon Salmo salar proposes that North American and southern European stocks (<62° N) move directly to feeding grounds off west Greenland, then overwinter in the Labrador Sea, whereas northern European stocks (>62° N) utilize the Norwegian Sea. An alternate model proposes that both North American and European stocks migrate in the North Atlantic Subpolar Gyre (NASpG) where S. salar enter the NASpG on their respective sides of the Atlantic, and travel counterclockwise within the NASpG until returning to natal rivers. A review of data accumulated during the last 50 years suggests a gyre model is most probable. Freshwater parr metamorphose into smolts which have morphological, physiological and behavioural adaptations of epipelagic, marine fishes. Former high‐seas fisheries were seasonally sequential and moved in the direction of NASpG currents, and catches were highest along the main axis of the NASpG. Marking and discrimination studies indicate mixed continental origin feeding aggregations on both sides of the Atlantic. Marked North American smolts were captured off Norway, the Faroe Islands, east and west Greenland, and adults tagged at the Faroes were recovered in Canadian rivers. Marked European smolts were recovered off Newfoundland and Labrador, west and east Greenland, and adults tagged in the Labrador Sea were captured in European rivers. High Caesium‐137 (¹³⁷Cs) levels in S. salar returning to a Quebec river suggested 62·3% had fed at or east of Iceland, whereas levels in 1 sea‐winter (SW) Atlantic Canada returnees indicated 24·7% had fed east of the Faroes. Lower levels of ¹³⁷ Cs in returning 1SW Irish fish suggest much of their growth occurred in the western Atlantic. These data suggest marine migration of S. salar follows a gyre model and is similar to other open‐ocean migrations of epipelagic fishes.     

Evidence for an autumn downstream migration and the subsequent estuarine residence of 0+ year juvenile Atlantic salmon Salmo salar L., in England

The use of passive integrated transponder technology facilitated the first observations of the autumn migration of 0+ year Atlantic salmon Salmo salar parr into tidal rearing habitats in the U.K. The quality of these habitats in relation to smolt production and the ecological significance of this alternative life-history strategy are presently not understood.     

Migratory behaviour of post-smolt Atlantic salmon during initial stages of seaward migration

The movements of 24 hatchery-reared Atlantic salmon Salmo salar smolts, with miniature acoustic transmitters (pingers) implanted surgically, were determined after release in the coastal waters of Passamaquoddy Bay (mean tide range 6 m), New Brunswick, Canada, to describe the first stages of seaward migration. Automated pinger detection at fixed sites, and pinger location and tracking by boat were used. Post-smolts left the release area rapidly, and the majority moved to open waters of the bay within several tidal cycles. Initially, post-smolts moved with a seaward orientation on ebb tides and held positions on flood tides. Their movements into open waters were diurnal, and the timing corresponded with the state of the tide during which they moved through a narrow channel. Post-smolts moved preferentially through this passageway with the aid of the tidal stream. Successful movements out through the channel occurred during ebb tides and any movements back in were during flood tides. Ground speed of fish moving through the channel was 4·2 body lengths s⁻¹ and faster than the tidal stream velocities in the channel. The relative velocity of fish swimming through the channel was 2 body lengths s⁻¹. Post-smolt movement was indicative of active, directed swimming with a reliance on ebb-tide transport for migration through a coastal area with strong tidal currents. Some post-smolts moved seaward directly with no apparent period of acclimation for the transfer to the marine environment, whereas others delayed their departure. These differences in behaviour were probably related to asynchrony in smolting when fish were released.     

Observations on the biology and ecology of the blue shark in the North-east Atlantic

Of 159 blue sharks Prionace glauca examined (59% female, 41% male) from oceanic waters SW of Britain and Ireland, all but two males were immature. Size-at-age and growth rate were similar to previous studies in the North Atlantic, while stomach contents included cephalopods, fish, cetaceans, and to a lesser degree, birds and crustaceans.     

Return migration of Atlantic salmon in the River Tana: the role of environmental factors

Multi-sea-winter Atlantic salmon (75–115 cm fork length L _F, 2–4-winter fish) were radio-tagged in the Tanafjord (700 N), Norway, in 1992–1993, and 130 fish entered the large subarctic River Tana (Teno). They entered the fresh water at any time of the tidal cycle but more so during the high and ebbing tides. No diel rhythm was detected in river entry under polar day conditions. There were no differences in the change of flow between days when salmon moved and when they did not, but during active migration increasing discharge was associated with increased swimming activity of salmon, especially later in the summer. Increasing air temperature was also associated with enhanced migration activity. Low river flow was associated with increasing delay in salmon passing the first riffle area of the river, 35 km from the sea.     

Migration success and migration time of gastrically radio-tagged v. PIT-tagged adult Atlantic salmon

Adult Atlantic salmon (55–116 cm total length) with PIT-tags ( n = 318) and both PIT-tags and gastric radio-tags ( n = 127) in a regulated river showed similar upstream migration success, yet radio-tagged fish migrated faster than fish with only PIT-tags. Regurgitation rate of radio-tags was 9%.     

Frequency of vateritic otoliths and potential consequences for marine survival in hatchery-reared Atlantic salmon

Otoliths are inner-ear structures of all teleost fish with functional importance for hearing and balance. The otoliths usually consist of aragonite, a polymorph of calcium carbonate, but may also take the form partly or entirely of vaterite, a different polymorph of calcium carbonate. Vateritic otoliths occur sporadically in wild fish, but with a higher frequency in hatchery-reared fish. Abnormal otoliths have direct consequences for the inner-ear functions of fish and may be a symptom of environmental stress. In this study, the authors assess the differences in the frequency of abnormal otoliths and degree of abnormality (% vaterite) for different groups of hatchery-reared Atlantic salmon (Salmo salar) smolt and adults. The groups differed in parental broodstock origin (number of generations in hatchery) and treatment temperature. Smolt from the same groups were also released to complete their ocean migration. The otoliths of the returning and recaptured adults were subsequently extracted to assess the difference in frequency and degree of abnormality between the adults and the smolt from corresponding groups. Return rate varied among groups (0.2%–2.6%). The frequency of vateritic otoliths was high (11.4%–64.4%) and differed among smolt groups. The lowest return rates corresponded with the highest frequency of abnormal otoliths for the groups, suggesting that abnormal otoliths may have negative consequences for marine survival. Furthermore, indications of an effect of fast growth on the formation of abnormal otoliths were found for only one of the experimental groups, and for none of the groups after correcting for Type 1 error. This contradicts previous reports, suggesting rapid growth as the main cause of abnormal otoliths. Adult return rates were generally low, but abnormal otoliths were common, with high coverage (% vaterite).     

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).     

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).