Associate Professor

Pathology and Anatomical Sciences




  • B.S. Biology, Wake Forest University (Summa Cum Laude), 1995
  • Ph.D. Ecology and Evolutionary Biology, Brown University, 2003

Academic Appointments

  • Research Associate, Department of Ecology and Evolutionary Biology, Brown University
  • NIH Ruth Kirschstein National Research Service Award Fellow, University of California, Riverside and Brown University

Website: Middleton Lab

Research Description

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.


  • Biomechanics and locomotion
  • Skeletal physiology and adaptation
  • Experimental evolution

Representative Publications

  1. Middleton, KM, SA Kelly, and T Garland, Jr. 2008. Selective breeding as a tool to probe skeletal response to high voluntary locomotor activity in mice. Integrative and Comparative Biology 48: 394–410.
  2. Middleton, KM, BD Goldstein, PR Guduru, JF Waters, SA Kelly, SM Swartz, and T Garland, Jr. 2010. Variation in within-bone stiffness measured by nanoindentation in mice bred for high levels of voluntary wheel running. Journal of Anatomy 216: 121–131.
  3. Garland, T Jr, SA Kelly, JL Malisch, EM Kolb, RM Hannon, BK Keeney, SL Van Cleave, KM Middleton. 2011. How to run far: multiple solutions and sex-specific responses to selective breeding for high voluntary activity levels. Proceedings of the Royal Society B 278: 574–581.
  4. Keeney, BK, TH Meek, KM Middleton, and T Garland, Jr. 2012. Sex differences in cannabinoid receptor-1 (CB1) pharmacology in mice selectively bred for high voluntary wheel-running behavior. Pharmacology, Biochemistry and Behavior 101: 528-537.
  5. Altshuler, DL, EM Quicazán-Rubio, PS Segre, KM Middleton. 2012. Wingbeat kinematics and motor control of yaw turns in Anna’s hummingbird (Calypte anna). Journal of Experimental Biology 215: 4070-4084.