State-dependent behavior and alternative behavioral strategies in brown trout (Salmo trutta L.) fry

Animals generally adjust their behavior in response to bodily state (e.g., size and energy reserves) to optimize energy intake in relation to mortality risk, weighing predation probability against the risk of starvation. Here, we investigated whether brown trout Salmo trutta adjust their behavior in relation to energetic status and body size during a major early-life selection bottleneck, when fast growth is important. Over two consecutive time periods (P1 and P2; 12 and 23 days, respectively), food availability was manipulated, using four different combinations of high (H) and low (L) rations (i.e., HH, HL, LH, and LL; first and second letter denoting ration during P1 and P2, respectively). Social effects were excluded through individual isolation. Following the treatment periods, fish in the HL treatment were on average 15-21 % more active than the other groups in a forced open-field test, but large within-treatment variation provided only weak statistical support for this effect. Furthermore, fish on L-ration during P2 tended to be more actively aggressive towards their mirror image than fish on H-ration. Body size was related to behavioral expression, with larger fish being more active and aggressive. Swimming activity and active aggression were positively correlated, forming a behavioral syndrome in the studied population. Based on these behavioral traits, we could also distinguish two behavioral clusters: one consisting of more active and aggressive individuals and the other consisting of less active and aggressive individuals. This indicates that brown trout fry adopt distinct behavioral strategies early in life. This paper provides information on the state-dependence of behavior in animals, in particular young brown trout. On the one hand, our data suggest a weak energetic state feedback where activity and aggression is increased as a response to short term food restriction. This suggests a limited scope for behavioral alterations in the face of starvation. On the other hand, body size is linked to higher activity and aggression, likely as a positive feedback between size and dominance. The experiment was carried out during the main population survival bottleneck, and the results indicate that growth is important during this stage, as 1) behavioral compensation to increase growth is limited, and 2) growth likely increases the competitive ability. However, our data also suggests that the population separates into two clusters, based on combined scores of activity and aggression (which are positively linked within individuals). Thus, apart from an active and aggressive strategy, there seems to be another more passive behavioral strategy.