Life in varying environments: experimental evidence for delayed effects of juvenile environment on adult life history.

1. The effects of environment experienced during early development on phenotype as an adult has started to gain vast amounts of interest in various taxa. Some evidence on long-term effects of juvenile environment is available, but replicated experimental studies in wild animals are still lacking. 2. Here we report the first replicated experiment in wild mammals which examines the long-term effects of juvenile and adult environments on individual fitness (reproduction, survival and health). The early development of bank vole (Myodes glareolus) individuals took place in either food-supplemented or un-supplemented outdoor enclosures. After the summer, adult individuals were reciprocally changed to either a similar or opposite resource environment to overwinter. 3. Adult environment had an overriding effect on reproductive success of females so that females overwintering in food-supplemented enclosures had a higher probability of breeding and advanced the initiation of breeding. However, the characteristics of their litters were determined by juvenile environment: females initially grown in food-supplemented conditions subsequently produced larger litters with bigger pups and a male-biased sex ratio. 4. In males, individuals growing in un-supplemented conditions had the highest survival irrespective of adult environment during winter, whereas in females, neither the juvenile nor adult environments affected their survival significantly. The physiological condition of voles in spring, as determined by haematological parameters, was also differentially affected by juvenile (plasma proteins and male testosterone) and adult (haematocrit) environments. 5. Our results suggest that (i) life-history trajectories of voles are not strictly specialized to a certain environment and (ii) the plastic life-history responses to present conditions can actually be caused by delayed effects of the juvenile environment. More generally, the results are important for understanding the mechanisms of delayed life-history effects as well as recognizing their population dynamic consequences.