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This hypothesis implies that social decision-making mechanisms localized in the central nervous system CNS are open to the influence of peripheral hormones that ultimately are under the control of the CNS through the hypothalamic-pituitary-gonadal axis. Therefore, two puzzling questions emerge at two different levels of biological analysis: 1 Why does the brain, which perceives the social environment and regulates androgen production in the gonad, need feedback information from the gonad to adjust its social decision-making processes?
In this paper, we will address these two questions using the integrative approach proposed by Niko Tinbergen, who proposed that a full understanding of behavior requires its analysis at both proximate physiology, ontogeny and ultimate ecology, evolution levels.
Although these four questions can be interpreted as culminating into the proximate-ultimate dichotomy of biological causation proposed by Mayr , Tinbergen's formulation clearly distinguishes cause from function and calls not the separateness of his questions, but rather for their integration when investigating a particular phenotype.
Only such an integrative approach would allow a truly comprehensive understanding of the behavior in question. Indeed, on one hand knowledge of the proximate mechanisms underlying a given behavior is crucial to understanding its costs, limits and evolutionary consequences, therefore highlighting the fact that proximate mechanisms contribute to the dynamics of selection.
On the other hand, knowledge on the ecological function and evolution of a given behavior will clarify how the proximate mechanisms underlying it evolved. Thus, reciprocal causation analysis of biological phenomena i. Following Tinbergen's footsteps, here we aim to integrate the study of function with the study of proximate mechanisms of the social modulation of androgens.