Hormones and phenotypic plasticity: Implications for the evolution of integrated adaptive phenotypes

It is generally accepted that taxa exhibit genetic variation in phenotypic plasticity, but many questions remain unanswered about how divergent plastic responses evolve under dissimilar ecological conditions. Hormones are signaling molecules that act as proximate mediators of phenotype expression by regulating a variety of cellular, physiological, and behavioral responses. Hormones not only change cellular and physiological states but also influence gene expression directly or indirectly, thereby linking environmental conditions to phenotypic development. Studying how hormonal pathways respond to environmental variation and how those responses differ between individuals, populations, and species can expand our understanding of the evolution of phenotypic plasticity. Here, we explore the ways that the study of hormone signaling is providing new insights into the underlying proximate bases for individual, population or species variation in plasticity. Using several studies as exemplars, we examine how a 'norm of reaction' approach can be used in investigations of hormone-mediated plasticity to inform the following: 1) how environmental cues affect the component hormones, receptors and enzymes that comprise any endocrine signaling pathway, 2) how genetic and epigenetic variation in endocrine-associated genes can generate variation in plasticity among these diverse components, and 3) how phenotypes mediated by the same hormone can be coupled and decoupled via independent plastic responses of signaling components across target tissues. Future studies that apply approaches such as reaction norms and network modeling to questions concerning how hormones link environmental stimuli to ecologically-relevant phenotypic responses should help unravel how phenotypic plasticity evolves.