Reformulation of Trivers–Willard hypothesis for parental investment

The Trivers-Willard hypothesis (TWH) plays a central role in understanding the optimal investment strategies to male and female offspring. Empirical studies of TWH, however, yielded conflicting results. Here, we present models to predict optimal comprehensive multi-element parental strategies composed of primary sex ratio, brood size, resource allocation among offspring, and the resultant secondary sex ratio. Our results reveal that the optimal strategy depends on sex differences in the shape of offspring fitness function rather than in fitness variance. Also, the slope of the tangent line (through the origin) to the offspring fitness function can be used to predict the preferred offspring sex. We also briefly discuss links between the model and the empirical research. This comprehensive reformulation of TWH will offer a thorough understanding of multi-element parental investment strategies beyond the classical TWH.