Physiological epistasis, ontogenetic conflict and natural selection on physiology and life history

Ontogenetic conflict arises when optima for alleles governingfitness variation differ between juveniles and adults or betweenadult sexes. Loci that govern development of alternative phenotypesin the sexes, hereafter termed morph-determining loci, mediatedevelopment through the endocrine system. Morphotypic selectionis defined to be multivariate selection favoring discrete alternativemorphotypes (e.g., optima). When the optimal combinations ofalleles for alternative morphs differ between the sexes, itgenerates conflicting selection pressure and thus ontogeneticconflict. Selection on morph alleles promotes ontogenetic conflictbecause it perturbs physiological epistasis that governs theexpression of male versus female traits. Expression of physiologicaltraits arises from homeostasis that maintains trait expressionwithin a normal range. The genetic basis of homeostasis is likelyto arise from interactions among several genes (e.g., geneticepistasis) or protein products (e.g., physiological epistasis).For example, endocrine regulation arises from interactions betweengondatropins, which are protein hormones produced by the hypothalamic-pituitaryglands, and steroid hormones, which are produced by the gonads(e.g., HPG axis). The side-blotched lizard system is discussedwith respect to physiological bases of ontogenetic conflict.We also describe a novel molecular marker strategy for uncoveringgenome-wide physiological epistasis in nature. Finally, ontogeneticconflict exerts selection on females to evolve mate selectionor cryptic choice that is reflected in different sires beingchosen for son versus daughter production. We describe how side-blotchedlizard females ameliorate ontogenetic conflict by cryptic choiceof male genotypes to produce sons versus daughters.