The influence of freshwater environment on muscle growth in seawater was investigated in an inbred population of farmed Atlantic salmon (Salmo salar L.). The offspring from a minimum of 64 families per group were incubated at either ambient temperature (ambient treatment) or in heated water (heated treatment). Growth was investigated using a mixed-effect statistical model with repeated measures, which included terms for treatment effect and random fish effects for individual growth rate (α) and the instantaneous growth rate per unit change in temperature (γ). Prior to seawater transfer, fish were heavier in the heated (61.6±1.0·g; N=298) than in the ambient (34.1±0.4·g; N=206) treatments, reflecting their greater growth opportunity: 4872·degree-days and 4281·degree-days, respectively. However, the subsequent growth rate of the heated group was lower, such that treatments had a similar body mass (3.7-3.9·kg) after approximately 450·days in seawater. The total crosssectional area of fast muscle and the number (FN) and size distribution of the fibres was determined in a subset of the fish. We tested the hypothesis that freshwater temperature regime affected the rate of recruitment and hypertrophy of muscle fibres. There were differences in FN between treatments and a significant age×treatment interaction but no significant cage effect (ANOVA). Cessation of fibre recruitment was identified by the absence of fibres of <10·µm diameter. The maximum fibre number was 22.4% more in the ambient (9.3×10 5 ±2.0×10 4 than in the heated (7.6×10 5 ±1.5×10 4 ) treatments (N=44 and 40 fish, respectively; P<0.001). For fish that had completed fibre recruitment, there was a significant correlation between FN and individual growth rate, explaining 35% of the total variation. The density of myogenic progenitor cells was quantified using an antibody to c-met and was approximately 2-fold higher in the ambient than in the heated group, equivalent to 2-3% of the total muscle nuclei. The number of myonuclei in isolated fibre segments showed a linear relationship with fibre diameter. On average, there were 20.6% more myonuclei in 200-µm-diameter fibres isolated from the ambient (3734·myonuclei·cm -1 ) than from the heated (3097·myonuclei·cm -1 ) treatments. The maximum fibre diameter was greater in heated than in ambient groups, whereas the age×treatment interaction was not significantly different (ANCOVA). There were also no consistent differences in the rate of hypertrophy of muscle fibres between treatments. It was concluded that freshwater temperature regime affected fibre number and the nuclear content of fast muscle in seawater but not the rate of fibre hypertrophy. The mechanisms and life history consequences of developmental plasticity in fibre number are discussed.
SUMMARYAtlantic salmon (Salmo salar L.) were fed to satiety and reared from ∼60 g to 5000 g at ambient seawater temperatures. The effect of photoperiod manipulation on muscle growth was investigated from the start of the first sea winter. Continuous light treatment in winter/spring (1 November to 18 June) improved growth performance in fish, resulting in a 30% increase in mean body mass relative to the ambient photoperiod fish by 12 August, but had no effect on sexual maturation. Significant increases in body mass in the continuous light groups were observed after 126 days (P<0.01). The number of fast muscle fibres per trunk cross-section was determined in a subset of the fish and was 28.5% higher in the continuous light(799×103) than the natural day length(644×103) groups after only 40 days, corresponding to the period of decreasing natural day length. Subsequent rates of fibre recruitment were similar between treatments. At the end of the fibre recruitment phase of growth (combined June and August samples), the maximum number of fast muscle fibres was 23% higher in fish from the cages receiving continuous light(881×103±32×103; N=19) than in the ambient photoperiod cages(717×103±15×103; N=20)(P<0.001). Continuous light treatment was associated with a shift in the distribution of fibre diameters, reflecting the altered patterns of fibre recruitment. However, the mean rate of fibre hypertrophy showed no consistent difference between treatments. There was a linear relationship between the myonuclear content of isolated single fibres and fibre diameter. On average, there were 27% more myonuclei in 150 μm-diameter fibres in the continuous light (3118 myonuclei cm-1) than the ambient photoperiod(2448 myonuclei cm-1) fish. After 40 days, continuous light treatment resulted in a transient increase in the density of myogenic progenitor cells, identified using a c-met antibody, to a level 70% above that of fish exposed to natural light. It is suggested that short days inhibited the proliferation of myogenic progenitor cells and that this was overcome by transferring fish to continuous light, causing an increase in the number of times the myogenic precursor cells divided and/or a decrease in cell cycle time. The net increase in myogenic progenitor cells resulted in proportional increases in the number and myonuclear content of fibres. The subsequent hypertrophy of these additional fibres can explain the delayed increase in body mass observed with continuous light treatment.
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