A.A.P. conceived the study. C.C.P. and A.A.P. supervised the project. S.G. and P.C. designed and implemented the statistical and bioinformatics analyses with contributions from C.C.P., J.K.P. and A.A.P. N.M.G. designed and executed the RNA-seq and GBS protocols with assistance from E.A. and J.D. C.C.P. performed the behavioral phenotyping with assistance from E.L. and Y.J.P. A.L. performed the muscle and bone phenotyping with input from D.A.B. C.L.A.-B. performed the BMD phenotyping. C.C.P., S.G., P.C. and A.A.P. wrote the manuscript, with input from all co-authors.