Watching the days go by: Asymmetric regulation of caterpillar development by changes in photoperiod

Many insects possess the plastic ability to either develop directly to adulthood, or enter diapause and postpone reproduction until the next year, depending on environmental cues (primarily photoperiod) that signal the amount of time remaining until the end of the growth season. These two alternative pathways often differ in co-adapted life-history traits, for example, with slower development and larger size in individuals headed for diapause. The developmental timing of these differences may be of adaptive importance: If traits diverge early, the potential for phenotypic differences between the pathways is greater, whereas if traits diverge late, the risk may be lower of expressing a maladaptive phenotype if the selective environment changes during development. Here, we explore the effects of changes in photoperiodic information during life on pupal diapause and associated life-history traits in the butterfly Pararge aegeria. We find that both pupal diapause and larval development rate are asymmetrically regulated: While exposure to long days late in life (regardless of earlier experiences) was sufficient to produce nondiapause development and accelerate larval development accordingly, more prolonged exposure to short days was required to induce diapause and slow down prediapause larval development. While the two developmental pathways diverged early in development, development rates could be partially reversed by altered environmental cues. Meanwhile, pathway differences in body size were more inflexible, despite emerging late in development. These results show how several traits may be shaped by the same environmental cue (photoperiod), but along subtly different ontogenies, into an integrated phenotype.