1. Much attention has been devoted to explaining the spatial distribution of foraging animals, but rather little to their temporal distribution (i.e. whether they are diurnal, nocturnal or crepuscular). Many animals face predictable diel cycles of food availability or predation risk, and so the approach of measuring the relative ratio of mortality risk to food gained (the ‘minimize μ/f’ rule) can be applied equally as well to different time periods of the day as to alternative food patches or habitats. 2. This method is used here to investigate the diel activity patterns of juvenile Atlantic salmon, which have previously been shown to become increasingly biased towards nocturnal activity in winter, hiding for much of the day in streambed refuges. Calculations based on published data show that nocturnal foraging in winter is far safer per unit of food obtained than is diurnal, despite greatly reduced food capture efficiency at night‐time light levels. 3. Using an automated activity monitoring system based on passive integrated transponder (PIT) tags, this study shows that winter diel activity patterns in salmon are dependent on food availability. A change in food density led to a parallel change in time spent in the refuge, but (as predicted by the μ/f rule) the effect was greatest at the time of day with the least favourable ratio of predation cost to feeding benefit. Thus an experimental increase in food availability led to a 16% reduction in time spent in nocturnal foraging but a 98% reduction in time spent foraging by day, with fish spending only 0·6% of the daylight hours out of the refuge at the highest food density. 4. However, brief daytime foraging bouts had a major impact on growth rates (presumably because feeding efficiency was much greater in daylight), especially when food was scarce. Daytime feeding was thus profitable in terms of rapid food acquisition but normally suboptimal in terms of risk of predation. 5. Daily activity patterns are therefore suggested to be the result of a complex trade‐off between growth and survival, which takes account of diel fluctuations in food availability, food capture efficiency and predation risk; individual variation in the extent of diurnal feeding in salmon may result from state‐dependent differences in the benefits of rapid feeding and growth.

The diel and seasonal activity patterns of salmonids are predominantly governed by the annual changes in photoperiod and temperature. In winter salmonids become increasingly nocturnal, hiding in refuges by day but emerging to feed at night. This behaviour may be linked to either one of the controlling influences mentioned above or to an inherent annual rhythm. Here we show that the previously described switch by Altantic salmon (Salmo salar L.) from predominantly diurnal to nocturnal activity in winter also occurs at other times of the year in response to low, "winter" temperatures; this is demonstrated both in laboratory experiments and by field observations in glacial rivers. This indicates that there is no underlying inherent annual rhythm to this behaviour nor any photoperiodic influence. Furthermore, in the laboratory experiment this temperature-dependent shift to nocturnalism was explained by a suppression of daytime activity rather than an increase in activity at night.

1. Most animals are active by day or by night, but not both; juvenile salmonids are unusual in that they switch from being predominantly diurnal for most of the year to being nocturnal in winter. They are visual foragers, and adaptations for high visual acuity at daytime light intensities are generally incompatible with sensitive night vision. Here we test whether juvenile Atlantic Salmon Salmo salar are able to maintain their efficiency of prey capture when switching between diurnal and nocturnal foraging. 2. By testing the ability of the fish to acquire drifting food items under a range of manipulated light intensities, we show that the foraging efficiency of juvenile salmon is high at light intensities down to those equivalent to dawn or dusk, but drops markedly at lower levels of illumination: even under the best night condition (full moon and clear sky), the feeding efficiency is only 35% of their diurnal efficiency, and fish will usually be feeding at less than 10% (whenever the moon is not full, skies are overcast or when in the shade of bankside trees). Fish were unable to feed on drifting prey when in complete darkness. 3. The ability of juvenile salmon to detect prey under different light intensities is similar to that of other planktivorous or drift‐feeding species of fish; they thus appear to have no special adaptations for nocturnal foraging. 4. While winter drift abundance is slightly higher by night than by day, the difference is not enough to compensate for the loss in foraging efficiency. We suggest that juvenile salmon can nonetheless switch to nocturnal foraging in winter because their food requirements are low, many individuals adopting a strategy in which intake is suppressed to the minimum that ensures survival.

Animal species have usually evolved to be active at a speci¢c time of the daily cycle, and so are either diurnal, nocturnal or crepuscular. However, we show here that the daily timing of activity in juvenile Atlantic salmon is related to the life-history strategy that they have adopted (i.e. the age at which they will migrate to the sea) and their current state (body size/relative nutritional state). Salmon can detect food more easily by day than by night, but the risk of predation is greater. Nocturnal foraging should generally be preferred, but the greater the need for growth, the greater should be the shift towards diurnal activity. In line with this prediction, all ¢sh were predominantly nocturnal, but salmon preparing to migrate to the sea, which would experience size-dependent mortality during the forthcoming migration, were more diurnal than ¢sh of the same age and size that were delaying migration for a further year. Moreover, the proportion of activity by day was negatively correlated with body size within the intending migrants. It has previously been shown that overwinter survival in ¢sh delaying migration is maximized not by growth but by minimizing exposure to predators. As predicted, daytime activity in these ¢sh was correlated with the prior rate of weight loss, ¢sh being more diurnal when their risk of starvation was greater. To our knowledge, these are the ¢rst quantitative demonstrations of state-dependent variation in the timing of daily activity.

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The activity patterns of animals, whether diurnal, crepuscular or nocturnal, are usually fixed endogenous rhythms, entrained by environmental Zeitgebers. Here we demonstrate that juvenile Atlantic salmon, Salmo salar, switch between diurnal and nocturnal foraging solely in response to environmental temperature, and independently of photoperiod and season. Above 10 °C juvenile Atlantic salmon fed predominantly during daylight, spending the night exposed in the water column but relatively quiescent. As temperature dropped below 10 °C they became increasingly nocturnal, hiding in refuges by day but emerging to feed at night. It has previously been shown that parallel physiological changes take place in the retinae of several species of salmonids : the quantity and composition of the visual pigments change so as to make the fish more dark adapted at low temperatures. As the fish were found to be far less aggressive by night than by day at all temperatures, the switch to nocturnal activity was also accompanied by a change in social structure. We suggest that this temperature-dependent strategy maximizes feeding efficiency in summer but reduces predation risk in winter.

Dispersal and growth were studied in Atlantic salmon, Salmo salar L., fry from a natural isolated redd. The distribution of fry leaving the redd was strongly peaked, 80% being caught within a 2‐week period. Early in the dispersal period, all fry leaving the redd had remnants of yolk sac and had not fed; by half way through the dispersal period, no fry had any visible yolk, but 35% still had empty stomachs. Fry leaving the redd during the first half of the dispersal period tended to settle in different first feeding sites than those dispersing later. Predation on fry by larger salmonids was frequent, especially during dispersal. Coefficients of variation for length, weight and condition factor increased significantly over the study period and for individually recognised fry, growth rates varied markedly. Thus, individual salmon fry differ in physical status on emergence from the redd and these differences are amplified during the first few weeks after emergence.