Abstract
The motor systems that produce behavioral movements are among the primary targets for the action of steroid hormones, including androgens. Androgens such as testosterone bind to androgen receptors (AR) to induce physiological changes in the size, strength, and energetic capacity of skeletal muscles, which can directly influence the performance of behaviors in which those muscles are used. Because tissues differentially express AR, resulting in tissue-specific sensitivity to androgens, AR expression may be a major target of selection for the evolution of behavior. Anolis lizards (i.e., anoles) provide a robust system for the study of androgen-regulated traits, including the behavioral traits that facilitate social display and locomotion. In this study, we examined six anole species that demonstrate significant variation in the behavioral use of the forelimbs to measure the proportion of myonuclei in the bicep muscles that express AR. Using phylogenetic comparative analyses, we found that species with a greater proportion of nuclei positive for AR expression in the biceps exhibited greater frequencies of locomotor movements and pushup displays. These results suggest that AR expression in skeletal muscles may influence the evolution of androgen-regulated behaviors in this group.



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Acknowledgements
We thank our many colleagues and student collaborators for their assistance in the collection of lizard behavioral data over five field seasons; Kathleen Foster for advice on muscle morphology; and Brittney Ivanov, Jonathan King, Jimmy Roberts, and Mark Brodl for support in the laboratory. All procedures were performed with the approval of Trinity University’s Animal Research Committee (protocols 042811-MJ1, 011415_MJ1, NSF_050213_MAJ3), the Ministerio de Medio Ambiente y Recursos Naturales, Dominican Republic, and Texas Parks & Wildlife (permit SPR-0310-045). Funding for this work was provided by the National Science Foundation (IOS 1257021 to MAJ), an HHMI-funded undergraduate student fellowship (to DJC) from Trinity University, The Explorer’s Club Youth Activity Fund (to DJC), and the Mind Science Foundation (to DJC).
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Johnson, M.A., Kircher, B.K. & Castro, D.J. The evolution of androgen receptor expression and behavior in Anolis lizard forelimb muscles. J Comp Physiol A 204, 71–79 (2018). https://doi.org/10.1007/s00359-017-1228-y
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DOI: https://doi.org/10.1007/s00359-017-1228-y


