Social status, access to food, and compensatory growth in juvenile Atlantic salmon

Juvenile Atlantic salmon Salmo salar subjected to three weeks of cooler temperatures were 8·5% smaller than controls at the end of the temperature manipulation, but had caught up in size 20 weeks later. The behavioural means is examined by which this catch-up or compensatory growth is achieved. While on average compensating fish did not spend more time feeding, dominant fish within each group gained more exclusive access to the feeding area during periods of catch-up growth. Therefore the extent to which compensatory growth could be achieved was dependent on both the social status of the individual and the dominants' ability to monopolize the food patch.     

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

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

Social status and growth rates as determinants of site attachment in juvenile Atlantic salmon

This paper describes an experimental study of the effects of food supply, growth rates and social interactions on homing by juvenile Atlantic salmon Salmo salar in response to displacement. Groups of five fish were housed in a section of an artificial stream and given either rations allowing maximum growth (the rich condition) or 0·1 of this amount (the poor condition); daily specific growth rates were significantly higher in the rich condition. After a 6-day settlement period, the fish were captured, displaced downstream and their movements recorded over the next 3 h. Prior to displacement, the fish showed a high degree of site fidelity and high levels of aggression. Dominant fish and those with stronger site attachment grew faster prior to displacement, these effects being independent. Following displacement, 24% of all fish returned to their previously favoured site and stayed there, 23% returned home initially, but subsequently moved on, 5% settled in a new site and 49% failed to move. The distribution of responses was identical for the rich and poor conditions, but fish that homed were dominant and had grown faster during the pre-displacement period.     

Shelter selection in juvenile Atlantic salmon,or why do salmon seek shelter in winter?

During winter, juvenile Atlantic salmon Salmo salar become nocturnal and seek refuge during the day in the stream bed gravel interstitial spaces. The function of this behaviour is unclear, but two major types of hypothesis have been proposed. One is that the fish are hiding from something (e.g. a predator) and the other is that the fish are seeking shelter from the water current. These hypotheses were tested by examining the selection of juvenile salmon for refuges that offered different degrees of concealment or shelter. The fish clearly preferred refuges that allowed them to hide (i.e. they were dark and opaque) but offered little shelter from the current. Therefore, it can be assumed that the primary function of this nocturnal behaviour during winter is most likely to hide from diurnal predators.     

The effects of initial length and body curvature on shrinkage of juvenile Atlantic salmon during freezing

Fork length was measured in two groups of salmon parr (32–139 mm, frozen in a straight posture and frozen in a curved posture) before (L₁) and after (L₂) freezing and thawing. All the fish shrank. The decrease in length was significantly greater in the curved fish than the straight fish. The absolute reduction in length (L₁–L₂) was related directly to L₁, whereas the percentage reduction in length [(L₁–L₂)/L₁× 100] was related inversely to L₁.     

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

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

The feeding behaviour of juvenile Atlantic salmon in relation to turbulent flow

The feeding behaviour of juvenile Atlantic salmon Salmo salar in the Sainte‐Marguerite River, Quebec, Canada, varied with the characteristics of turbulent flow. Simulations indicated that juveniles would decrease their swimming costs during attacks by 19·8% in low and by 31·1% in high turbulent conditions by initiating movements in low‐speed flow events. The real swimming costs did not differ from the swimming costs estimated for a situation where fish initiate their movements at randomly selected flow velocities. The juvenile Atlantic salmon did not seem to prefer low‐speed flow events when initiating their movements. The proportion of time used for movements by fish decreased with an increase in the mean and the s . d . of the flow velocity.     

Seasonal growth of wild Atlantic salmon juveniles and implications on age at smoltification

The seasonal growth trajectories of wild Atlantic salmon Salmo salar juveniles by age group within the Margaree River, Canada, are described. Circuli counts from scales were used to infer growth rates at different ages and these were used to predict the proportions of age 2‐ and 3‐year old smolts from different portions of the watershed. In the wild Atlantic salmon juveniles from the Margaree River, there was no bimodality in fork length frequencies and no 1 year old smolts were produced. Water temperature differences during the growing season were insufficient to explain the differences in growth rates and size at age among the sites sampled. There was a positive association between the growth rate in the first year and the subsequent age at smoltification. In the Margaree River, differences in tributary specific growth rates and size at age were expected to produce important differences in the relative ages at smoltification.     

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.     

Compensatory sea growth of male Atlantic salmon, Salmo salar L., which previously mature as parr

The proportion of mature male parr in 11 families of Atlantic salmon, Sulrno sulur, reared under similar conditions in fresh water varied from 0–43%. The mature males were smaller than their siblings in December as 1 + and in late March. After individual tagging and transfer to a sea cage in early April. the previously mature males grew faster than previously immature salmon during the next 6 months. This compensatory growth resulted in almost equal size between the two groups. The results are discussed in relation to the different life strategies of salmon.     

Size and age of maturity of Atlantic salmon correlate with the North Atlantic Oscillation Index (NAOI)

Sea‐age at maturity of Atlantic salmon Salmo salar decreased with increasing values of the seasonal NAOI from February to April. Body mass increment from smolts to adults of one‐sea‐winter Atlantic salmon increased with increasing NAOI in May at the time when the juveniles moved to sea.     

Competition for shelter among over‐wintering signal crayfish and juvenile Atlantic salmon

Three separate effects on refuge use by signal crayfish Pacifastacus leniusculus and Atlantic salmon Salmo salar were examined: (1) the effect on Atlantic salmon of an addition of signal crayfish (doubling the total number of animals), (2) the effect on signal crayfish of an addition of Atlantic salmon and (3) intraspecific compared with interspecific competition, compared by holding total density of animals constant and varying the proportion of signal crayfish and Atlantic salmon in trials. Observations were made during winter, when both species are nocturnal. The proportion of Atlantic salmon sheltering was significantly lower in the presence than in the absence of signal crayfish when the interspecific treatment (Atlantic salmon plus signal crayfish) effected a doubling in density compared to the intraspecific treatment (Atlantic salmon alone). The proportion of signal crayfish sheltering was independent of the presence of Atlantic salmon. When total density was constant, the proportion of Atlantic salmon sheltering was significantly higher in intraspecific (52·8%) than interspecific trials (27·3%). Atlantic salmon out of shelter during the day in winter are believed to be very vulnerable to predators and the capacity for fish to share shelters with one another is known to be very low. Therefore, competition from crayfish for winter shelters may lead to detrimental effects on Atlantic salmon populations.     

Differences in growth between maturing and non-maturing male Atlantic salmon, Salmo salar L., parr

Sibling male Atlantic salmon parr that matured tended to be the larger fish in January, but their monthly specific growth rates between January and July did not differ from those of non-maturing fish. Maturing fish had lower condition factors in March, but greater increases in condition factor during April, exceeding those of non-maturing males by May. In maturing males, feeding rates between July and September, and specific growth rates in August and September, were lower than those of immature fish. Consequently, the mean size of immatures equalled or exceeded that of maturing males by October. Maturation rates were strongly correlated with increases in mean condition factor only during April.     

Individual space use strategies of wild juvenile Atlantic salmon

Movements of 60 stream-dwelling wild Atlantic salmon Salmo salar (97–118 mm), each tagged with a passive integrated transponder, were monitored during four trials in an enclosed section (24 m long, 45·1 m² total area) of a stream at a range of densities (four, eight, 16 and 32 fish per enclosure). Patterns of space use differed markedly between individuals, with 80% of fish establishing home ranges within 8 days of introduction to the enclosure (settlers) and the remainder continuing to move throughout the length of the enclosure (non-settlers). Although aggressive interactions were quite frequent and dominant fish were observed chasing subordinates, there was considerable overlap of home ranges of settlers at all densities; this was the case even at lower densities at which only a fraction of the enclosure was used by the fish. Thus, rather than adopting fixed territories, the salmon showed a high level of space sharing. Individual fish used the same local area in different ways, ranging from highly localized feeding on drifting food items to a wider-ranging strategy of specialising on benthic food. Among the fish that settled absolute growth rates were inversely related to body size, and at high densities fish lost weight. These results suggest that space use in wild juvenile salmon is more complex than a mosaic of territories, that salmon demonstrate significant variability in individual space use patterns, and that large fish may suffer disproportionately when populations are at the carrying capacity of their environment.     

Effects of life history variation on size and growth in stream-dwelling Atlantic salmon

A large size variation amongst life histories for stream-dwelling Atlantic salmon Salmo salar was found and the relative effect of life histories on size varied over time. As early as December (age 0+ years), fish that later smolted at age 2+ years were significantly larger than fish that did not smolt at age 2+ years. In contrast, there were no mass differences at age 0+ years between fish that would mature or not at age 1+ years (October). The mass differences between smolts and non-smolts persisted until smolting, and differences between mature and immature fish first appeared in May (age 1+ years). Following September (age 1+ years), there was also a significant interaction between smolting and maturity. Previously mature and immature age 2+ year smolts were not significantly different in size, but immature age 2+ year non-smolts were much lighter than mature age 2+ year non-smolts. Based on mass differences, the apparent 'decision' to smolt occurred c . 5 months before (winter, age 0+ years) the decision to mature (late spring, age 1+ years). In addition to strong seasonal growth variation, sizes of freshwater Atlantic salmon were largely structured by the complex interaction between smolt-age and maturity.     

Patterns of growth and smolting in autumn migrants from a Scottish population of Atlantic salmon, Salmo salar L.

In late November 1990 salmon parr, Salmo salar L., from the Girnock Burn in northern Scotland were either caught on their feeding territories (n=25) or trapped during downstream migration (n= 18). They were then housed in a laboratory rearing tank and their food intake and growth rates were tracked, until their smolting status was ascertained in the following May. Female fish were predominant in both groups; although the range of ages was the same, the total age of migrants was 2+ while that of residents was 1+. In November, compared to resident fish of the same year class, migrants were larger, heavier and in better condition. Although growth rates dropped during the winter in both groups before increasing in spring, migrants ate more and consistently grew faster than residents. In seawater tolerance tests conducted in May, more residents than migrants failed to adapt. These results confirm the suggestion that autumn migrants smolt in the following spring and suggest that they represent the faster-growing component of their cohort.     

The influence of life-history strategy on lipid metabolism in overwintering juvenile Atlantic salmon

From November to May, the lipid mass in the viscera and carcass of juvenile Atlantic salmon Salmo salar that were undergoing smolt transformation prior to seaward migration ('early migrants') were significantly greater than those of their siblings that would delay migration for at least a further year. During winter (November-February), the depletion of lipid associated with the viscera was significantly greater in early migrants, whilst lipid depletion in the remaining carcass was greater in delayed migrants. Early migrants continued to deplete both lipid compartments in spring (February-May), whereas delayed migrants depleted visceral lipid but replenished carcass lipid over the same period. Fatty acid accumulation rates (a measure of storage lipid synthesis rates) were two to six times greater in visceral than in carcass lipid throughout the study, suggesting that lipid turnover is much more rapid in the viscera. There were no differences in fatty acid accumulation rates between migrant groups in November, despite the much lower food consumption rate of delayed migrants at that time, suggesting that these fish allocated a larger proportion of their nutritional resources to lipid synthesis. In the carcass lipid of early migrants, and in both the visceral and carcass lipid of delayed migrants, the fatty acid accumulation rate was negatively correlated with lipid mass. Fatty acid accumulation rates increased from November to February in both visceral and carcass lipid in the two migrant groups. The fatty acid accumulation rate in carcass lipid was significantly higher in delayed migrants than in early migrants in February, but not in May. These results support the hypothesis that life history strategies involving rapid growth will result in a relatively low allocation of resources to lipid reserves.