Dr. Middleton’s research is focused on the integrative biology and evolution of vertebrate “locomotor tissues” including not only bones and muscles but also feathers and wing membranes. The interactions among these tissues are complex and occur at multiple anatomical levels, from the tissue to the structure as a whole. His lab integrates studies of locomotor tissues to understand how physiology, function, and evolution have shaped their structure—from microscale to macroscale—and how these changes are translated into locomotor differences among vertebrates.