The evolution of perennially enlarged breasts in women: a critical review and a novel hypothesis.

The possession of permanent, adipose breasts in women is a uniquely human trait that develops during puberty, well in advance of the first pregnancy. The adaptive role and developmental pattern of this breast morphology, unusual among primates, remains an unresolved conundrum. The evolutionary origins of this trait have been the focus of many hypotheses, which variously suggest that breasts are a product of sexual selection or of natural selection due to their putative role in assisting in nursing or as a thermoregulatory organ. Alternative hypotheses assume that permanent breasts are a by-product of other evolutionary changes. We review and evaluate these hypotheses in the light of recent literature on breast morphology, physiology, phylogeny, ontogeny, sex differences, and genetics in order to highlight their strengths and flaws and to propose a coherent perspective and a new hypothesis on the evolutionary origins of perennially enlarged breasts in women. We propose that breasts appeared as early as Homo ergaster, originally as a by-product of other coincident evolutionary processes of adaptive significance. These included an increase in subcutaneous fat tissue (SFT) in response to the demands of thermoregulatory and energy storage, and of the ontogenetic development of the evolving brain. An increase in SFT triggered an increase in oestradiol levels (E2). An increase in meat in the diet of early Homo allowed for further hormonal changes, such as greater dehydroepiandrosterone (DHEA/S) synthesis, which were crucial for brain evolution. DHEA/S is also easily converted to E2 in E2-sensitive body parts, such as breasts and gluteofemoral regions, causing fat accumulation in these regions, enabling the evolution of perennially enlarged breasts. Furthermore, it is also plausible that after enlarged breasts appeared, they were co-opted for other functions, such as attracting mates and indicating biological condition. Finally, we argue that the multifold adaptive benefits of SFT increase and hormonal changes outweighed the possible costs of perennially enlarged breasts, enabling their further development.