Much can be gained from the comprehensive study of a biological system. Based on what is known as Mayr's proximate-ultimate causation and the subsequent expansion to Tinbergen's four questions, biological traits can be understood by taking into account different approximations that try to explain mechanisms, development, adaptive significance or phylogeny. These, in principle, separate areas, can be integrated crossing boundaries, but bearing in mind that answers to one question would not explain a different query. Studies of sperm biology have, until now, not benefited much from this framework and potential integration. Proximate causes (particularly mechanisms) have been the subject of interest for reproductive biologists, and evolutionary explanations have been the domain of behavioural ecologists with interest in adaptive significance of traits in the context of post-copulatory sexual selection. This review will summarize opportunities for research in the different areas, focusing on sperm preparation for fertilization and suggesting possible integration within and between proximate and evolutionary studies.
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