The expensive son hypothesis

In its initial form, the expensive son hypothesis postulates that sons from male-biased sexually dimorphic species require more food during growth than daughters, which ultimately incur fitness costs for mothers predominantly producing and rearing sons. We first dissect the evolutionary framework in which the expensive son hypothesis is rooted, and we provide a critical reappraisal of its differences from other evolutionary theories proposed in the field of sex allocation. Then, we synthesize the current (and absence of) support for the costs of producing and rearing sons on maternal fitness components (future reproduction and survival). Regarding the consequences in terms of future reproduction, we highlight that species with pronounced sexual size dimorphism display a higher cost of sons than of daughters on subsequent reproductive performance, at least in mammals. However, in most studies, the relative fitness costs of producing and rearing sons and daughters can be due to sex-biased maternal allocation strategies rather than differences in energetic demands of offspring, which constitutes an alternative mechanism to the expensive son hypothesis stricto sensu. We observe that empirical studies investigating the differential costs of sons and daughters on maternal survival in non-human animals remain rare, especially for long-term survival. Indeed, most studies have investigated the influence of offspring sex (or litter sex ratio) at year T on survival at year T + 1, and they rarely provide a support to the expensive son hypothesis. On the contrary, in humans, most studies have focused on the relationship between proportion of sons and maternal lifespan, but these results are inconsistent. Our study highlights new avenues for future research that should provide a comprehensive view of the expensive son hypothesis, by notably disentangling the effects of offspring behaviour from the effect of sex-specific maternal allocation. Moreover, we emphasize that future studies should also embrace the mechanistic side of the expensive son hypothesis, largely neglected so far, by deciphering the physiological pathways linking son's production to maternal health and fitness.