Influence of habitat structure and fish density on Atlantic salmon Salmo salar L. territorial behaviour

A laboratory stream experiment was performed to test the influence of young‐of‐ the‐year Atlantic salmon Salmo Salar density on habitat selection ( i.e . pools and riffles) and the combined effect of habitat and density on territoriality. Pools were preferred at low and medium fish density. At higher density, the saturation of the riffles led to an equal fish repartition in both habitats. Aggressive interactions increased according to fish density, and the level of aggression was significantly higher in the pools than in the riffles.     

Effects of discharge and local density on the growth of juvenile Atlantic salmon Salmo salar

The study explored the combined effects of density, physical habitat and different discharge levels on the growth of juvenile Atlantic salmon Salmo salar in artificial streams, by manipulating flow during both summer and winter conditions. Growth was high during all four summer trials and increased linearly with discharge and mean velocity. Differences in fish densities (fish m^?3) due to differences in stream volume explained a similar proportion of the variation in mean growth among discharge treatments. Within streams, the fish aggregated in areas of larger sediment size, where shelters were probably abundant, while growth decreased with increasing densities. Fish appeared to favour the availability of shelter over maximization of growth. Mean growth was negative during all winter trials and did not vary among discharge treatments. These results suggest that increased fish densities are a major cause of reduced summer growth at low discharge, and that habitat‐mediated density differences explain the majority of the growth variation across habitat conditions both during summer and winter.     

Movement rhythms in juvenile Atlantic salmon, Salmo salar L.

Nocturnal downstrean migration of juvenile Atlantic salmon is usually interpreted as increased locomotor activity. The frequency of downstream passages of 0–1 + salmon in an endless stream channel was greater by night than by day in both smoking and non-smolting fish in autumn and spring. Movement increased at dusk, and decreased after dawn. Mature male 1 + fish moved slightly less than immatures in October, but significantly more in November. Total movement frequency was lower at full moon than at other moon phases, and movement was reduced when the moon was up. Under turbid conditions by day, the threshold water velocity inducing nett downstream movement was 8.2 cm s⁻¹, and the relative velocity of fish swimming downstream was never more than one third that of fish holding station at the normal maximal flow of 25–30 cm s⁻¹.
At the end of their first growing season in October, fish which had been offered food continuously through 24 h did not differ in size from those fed by day only, but the latter were significantly larger than those offered food only at night.
We conclude that: (1) the fish fed actively by day, and not by night; (2) station-holding represented activity, and downstream nocturnal movement represented relative inactivity (displacement) which occurred on loss of visual orientation, hence migration resulted from reduced activity; (3) lack of displacement in early autumn has adaptive value for maturing fish, but not for non-spawners.     

Prior residence, territory quality and life-history strategies in juvenile Atlantic salmon (Salmo salar L.)

Three groups of juvenile salmon were introduced sequentially into an artificial stream to investigate the effects of prior residence on behaviour and territory choice. Almost half of the first group obtained and defended distinct territories, the other half being constrained to an area approximately the size of one large territory. All of the fish in the subsequent groups, bar one, were also constrained to the same site. Since the fish were of similar size, prior residence alone seemed to influence which individuals obtained territories. However, within the first group, the fish that obtained territories were larger and more aggressive. The territorial fish did not appear to choose the most profitable territories, although they had the greatest opportunity to do so. Since juvenile salmon emerge from their gravel nests fairly synchronously, a time constraint on site sampling is hypothesised: there may be a risk in taking time to sample sites, since these same sites may become occupied with conspecifics. However, fish with territories fed at faster rates than non-territorial fish, possibly because of reduced competition for prey items. Consequently, fish from the first group (containing most of the territorial fish) grew faster than the other two groups. Moreover, most of the territory holders, but only one of the non-territorial fish, reached the threshold size that increases their probability of smolting the following year. This suggests that ability to obtain a defensible territory, primarily through prior residence, influences the age at which juvenile salmon can migrate to sea.     

Biochemical genetics of the Atlantic salmon Salmo salar L.

In recent years, blood group and protein polymorphisms in Atlantic salmon have been investigated extensively with a view, primarily, to their use in identifying individuals of different spawning populations present in high seas fisheries. Erythrocyte antigens, haemoglobins, serum proteins and various tissue enzymes—mainly esterases and de-hydrogenases—have been studied by electrophoretic and immunological techniques. These studies are reviewed here for the first time.
Many of the protein systems exhibit multiple components and this fact, together with cytological evidence, indicates the occurrence of tetraploidy in the course of Salmonid evolution. The significance of a tetraploid origin in the evolution and ecological adaptation of Salmonids is discussed briefly.
Some protein systems studied exhibit phylogenetic variation, and analyses of phenotype ratios and allele frequencies indicate that the populations of different river systems are genetically distinct. Allele frequencies have not yet been shown to be stable from generation to generation however, and some of the factors likely to affect allele frequencies are discussed.
Different spawning populations can not be identified in high seas fisheries using these protein characters, although it may be possible to identify the continent of origin (N. America or Europe) of some individuals. Indeed, it has been proposed independently by two groups that North American and European populations of Atlantic salmon be assigned to different sub-species viz. S. s. americanus and S. s. europaeus respectively. The contradictory evidence on which these taxa are proposed is discussed, together with the evidence for other population groups proposed in the European part of the salmon's range. The possible role and future direction of studies on the biochemical genetics of salmon are outlined.     

Influence of photoperiod on growth changes in juvenile Atlantic salmon, Salmo salar L.

During their first 6 months sibling Atlantic salmon parr, Salmo salar L., grew larger under constant light than under natural photoperiod or simulated natural photoperiod (control). When rate of change of photoperiod was accelerated after midsummer, ×2, ×3 and ×4, there were no growth differences between the three groups, but all were smaller than the control population. Under constant autumn photoperiod of 8 h light: 16 h dark growth was less than under all other experimental photoperiod conditions. Mean length was directly correlated with total hours of daylight experienced, excluding those fish kept under constant light. Fish reared from first feeding under photoperiod regimes delayed 6 and 9 months out of phase with the natural light cycle were smaller than the controls, whereas those under a regime 3 months out of phase did not differ from the controls. The clear segregation of modal length groups within the 3, 6 and 9 months out-of-phase populations occurred 1, 4 and 4 months, respectively, after the segregation in the control group. Under constant light, and under constant 12 h light: 12 h dark (12 LD), the segregation was delayed 3 and 4 months, respectively. The proportion of the population which maintained growth (upper modal group) was significantly less in the 9 and 6 months out-of-phase and 12 LD groups (39, 40 and 42%, respectively) than in the other three groups (82.5-85%). Acceleration of photoperiod change also resulted in decreased growth. The results support a model of salmon development in which, 2-3 months after first feeding, growth is maintained if feeding opportunities at that time are above a threshold level, and in which this critical timing is influenced by photoperiod. It is suggested that the delays reflect a synchronizing effect of photoperiod on an endogenous rhythm of appetite and growth. The differences in upper modal group proportions observed in the present experiments, reflect the relative feeding opportunities available at the critical period in July-August.     

The production of gynogenetic Atlantic salmon,Salmo salar L.

The production of Atlantic salmon gynogenomes by the combined use of a novel method for sperm irradiation and differently timed high hydrostatic pressure shocks is described. Sperm solutions were exposed to UV irradiation in a temperature-controlled flow-through device. Eggs fertilised with such sperm were exposed to shocks of 9500 psi at 30 min or approximately 7 h after fertilisation in order to produce meiotic and mitotic gynogenomes respectively. Yields of meiotic gynogenomes were generally high (up to 95%); those of mitotic gynogenomes were lower (range 2–20%). Analyses of the offspring by ploidy status and fingerprinting confirmed their gynogenetic origin. Small numbers of mitotic gynogenetic fish were grown on for 2 years in fresh and salt water. S1/S2 ratios were lower in gynogenetic fish and mean age at maturity was greater. Of the presumptive gynogenetic fish subjected to destructive sampling (n = 87) all were female.     

Periodicity in Atlantic salmon Salmo salar L. smolt migration

Behavioural and physiological mechanisms postulated for the control of downstream migration of Atlantic salmon smolts are reviewed briefly, and some new evidence is presented for their refusal to undergo sustained swimming. Although these mechanisms imply passive displacement as the primary means of emigration, it is likely that active components must also exist as the rates of travel of smolts through loch systems are only slightly slower than those recorded for river systems. The timing of these movements within 24 h periods is reviewed and it is shown that the predominantly nocturnal emigration pattern is evident on occasions in alevin, fry and parr stages also. Thus at migration the diel periodicity probably represents a seasonal locomotor rhythm which, under changed behavioural and physiological circumstances, results in downstream displacement.     

Liver ultrastructure of juvenile Atlantic salmon (Salmo salar)

Light and transmission electron microscopy of the liver of juvenile Atlantic salmon (Salmo salar) reveals a tubular arrangement of parenchymal cells, with biliary passages typically located at the center of tubules. Hepatocytes generally contain a single nucleus surrounded by a cuff of rough endoplasmic reticulum (RER), with many round to elongate mitochondria associated with the perinuclear RER. Whereas glycogen deposits are common and usually lie at the cell periphery, parenchymal cells seldom contain lipid droplets. Golgi complexes and heterogeneous dense bodies also occur in many hepatocytes, often in close proximity to bile canaliculi. Numerous microvilli from hepatocytes extend into the subendothelial space of Disse, which is also the location of stellate fat-storing cells. Interhepatocytic macrophages, sometimes containing prominent phagolysosomes and residual bodies, are common in the liver. The intrahepatic biliary system consists of intercellular canaliculi, bile pre-ductules, ductules, and ducts. In contrast to some other teleosts, the liver of the Atlantic salmon contains no intracellular bile canaliculi or Kupffer cells. The hepatic endothelium, arterioles, and perivenous regions are also described.     

Otolith shape discriminates between juvenile Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L.

Otoliths of Atlantic salmon, Salmo salar L., are more slender than the otoliths of brown trout, Salmo trutta L. Discriminant analysis on otolith measurements of juvenile Atlantic salmon and brown trout from four river systems revealed a discriminant function which distinguished more than 94% of the cases. This function was tested by using data from a fifth river with cohabiting Atlantic salmon and brown trout: all Atlantic salmon and 91 % of the brown trout were correctly classified.     

Awareness reactions and avoidance responses to sound in juvenile Atlantic salmon, Salmo salar L.

The possibility of using intense sound as a deterrent for juvenile Atlantic salmon ( Sulmo salar L. ) was studied by recording both physiological awareness reactions in an acoustic tube and behavioural avoidance responses in a pool. The measured awareness reactions consisted of decreased heart rate and breathing movements. Three criteria for the awareness reaction were used to compare the effect of different frequencies between 5 and 150 Hz: (i) threshold for spontaneous awareness reactions relative to the hearing thresholds, (ii) magnitude of the change in heart rate, (iii) degree of habituation to sound. After these criteria the lowest frequencies (5–10 Hz) were most effective in eliciting an awareness reaction from the fish. The responses of freely swimming fish to 10 and 150 Hz sounds were studied in an artificial pool. Juvenile salmon showed avoidance responses to 10 Hz stimulation at intensities 10–15 dB above the threshold for spontaneous awareness reactions measured in the acoustic tube. The 150 Hz sound failed to evoke avoidance responses, even at a level 30 dB above the threshold for spontaneous awareness reactions.     

Predicting microhabitat selection in juvenile Atlantic salmon Salmo salar by the use of logistic regression and classification trees

1. We compared the capacity of logistic regression (LR) and classification tree (CT) models to predict microhabitat use and the summer distribution of juvenile Atlantic salmon, Salmo salar, in two reaches of a small stream in eastern Quebec. 2. The models predicted the presence or absence of salmon at a location on the basis of habitat features (depth, current velocity, presence of instream and overhead cover, substratum particle size, and distance to stream bank) measured at that location. Models were validated by means of crossover field tests evaluating the performance of models developed for one reach (calibration trials) when applied to the other reach (validation trials). Model performance was evaluated with regard to accuracy, generality and ease of use and interpretation. Prediction maps based on habitat features were also built to compare the observed position of fish with those predicted by LR and CT models. 3. The spatial distribution of active fish differed markedly from that of resting fish, apparently as a result of the selection for water greater than about 30 cm depth by active fish and for the presence of rocky cover by resting fish. 4. All models made accurate predictions, validated by crossover trials. For both LR and CT models, the prediction maps reflected well the actual fish distributions. However, CT models were easier to build and interpret than LR models. CT models also had less variable performance and a smaller decline in predictive capability in crossover trials (for fish at rest), suggesting that they may be more transferable than LR models.     

Behavioural influences on life-history variation in juvenile Atlantic salmon,Salmo salar

Synopsis Juvenile Atlantic salmon emigrate from the river in a given year as a consequence of a physiological decision influencing appetite and growth during the previous summer. The direction of the decision depends on developmental performance exceeding a genetically determined threshold at that time, and that performance is governed by environmental opportunity. The animal's foraging efficiency (ability to avoid predators, and to compete) determines how well that opportunity is used. Those fish which maintained relatively high growth after July preferred higher stream flows, and were more likely to hide than to flee from a predator. Predator vigilance reduced intake, and ability to discriminate edible particles. Early competitive ability depended on fierceness rather than size, and ultimate large size was a consequence of dominant status. Initial status among high ranking individuals (but not among low ones) predicted the likelihood of maintaining growth in late summer. Size by July was the better predictor for low ranked fish. The influence of high status on life-history variation depends on how much it suppresses the growth of those lower in the hierarchy. Invited review     

Effect of water temperature and density of juvenile salmonids on growth of young-of-the-year Atlantic salmon Salmo salar

A von Bertalanffy growth model for young-of the-year Atlantic salmon Salmo salar in a small French coastal stream was fitted using water temperatures and densities of juvenile salmonids (S. salar and brown trout Salmo trutta) as covariates influencing daily growth rate. The Bayesian framework was used as a template to integrate prior information from external data sets. The relative influence of the covariates on parr growth was quantified and results showed that growth of S. salar juveniles depended on both water temperatures and densities, but that most of the spatiotemporal variability of growth resulted from local spatiotemporal variations of 0+ age salmonid (S. salar and S. trutta) densities. Further analysis revealed that the fluctuations in young-of-the-year salmonid densities are likely to dominate the effects of potential future warming of water temperature due to climate change. It is concluded that factors that could affect salmonid densities might well have a greater effect on S. salar population dynamics than factors influencing water temperatures.     

Observations on the gall bladder of juvenile Atlantic salmon Salmo salar L., in relation to feeding

Measurements of gut contents and changes in the volume and colour of bile in the gall bladder of juvenile Atlantic salmon are described for a variety of feeding regimes. The gall bladders of actively feeding fish are virtually empty of bile. When deprived of food, the weight of the gall bladder increases within 6 h and reaches a value equivalent to approximately 20% of the liver weight within 36 h. Bile is pale straw, green and blue in colour after 1, 4 and 6 days of starvation respectively. Stored bile is released from the gall bladder in response to food entering the anterior hind gut.     

Hyperplasia and hypertrophy in the growth of skeletal muscle in juvenile Atlantic salmon, Salmo salar L.

Both red and white muscle fibre numbers in juvenile Atlantic salmon increased gradually with fish length throughout the freshwater growth period. Mean fibre area increased as fish grew to 6.5 cm f.l. , but thereafter was unrelated to fish length. Hyperplasia was most obvious when fish were growing fastest, and was the dominant growth process in fish over 6.5 cm f.l. Hypertrophy was most important when growth was slow, as in autumn and winter.
Mean white fibre area was significantly smaller in deep muscle than at medial and superficial sites. Total cross-sectional area of red, white and total trunk muscle increased with fish length. The ratio of red: white cross-sectional area increased with fish length to a plateau at about 10% after 6.5 cm f.l.     

The role of food particle size in the growth of juvenile Atlantic salmon (Salmo salar L.)

The effect of particle size of a commercial pelleted feed on the growth of Atlantic salmon from first feeding alevins to first and second year smolts was investigated using 20-day feeding experiments. Ten experiments were performed over a two year period, each comprising six groups of fish separately selected from stock populations. Each group was presented with one of six sizes of food particle ranging from larger than the respective mean mouth breadth (100% feed size), through 50%, 25%, 12·5%, 6·25% to 3·125%. Experiments were performed in six radial flow/circumferential drain tanks under ambient photoperiod and water temperature. Growth rate was found to be closely related to feed size. Maximum growth in each case was shown only on one size of particle; larger and smaller sizes resulted in reduced growth. The particle size for maximum growth increased in direct proportion to fish length. Fish from 4·2 to 20·3 cm in length showed maximum growth on particle diameters 0·022 to 0·026 × fish fork length (PFR). First feeding alevins were found to show comparable growth rate on particle diameters 0·0115 to 0·090 PFR. Some seasonal variation in growth response was indicated. The results are discussed in relation to developmental and seasonal effects. Possible factors affecting the energetics of prey size related differential growth are discussed.     

Thermal habitat of adult Atlantic salmon Salmo salar in a warming ocean

The year-round thermal habitat at sea for adult Atlantic salmon Salmo salar (n = 49) from northern Norway was investigated using archival tags over a 10 year study period. During their ocean feeding migration, the fish spent 90% of the time in waters with temperatures from 1.6–8.4°C. Daily mean temperatures ranged from −0.5 to 12.9°C, with daily temperature variation up to 9.6°C. Fish experienced the coldest water during winter (November–March) and the greatest thermal range during the first summer at sea (July–August). Trends in sea-surface temperatures influenced the thermal habitat of salmon during late summer and autumn (August–October), with fish experiencing warmer temperatures in warmer years. This pattern was absent during winter (November–March), when daily mean temperatures ranged from 3.4–5.0°C, in both colder and warmer years. The observations of a constant thermal habitat during winter in both warmer and colder years, may suggest that the ocean distribution of salmon is flexible and that individual migration routes could shift as a response to spatiotemporal alterations of favourable prey fields and ocean temperatures.