Kevin Middleton, PhD

Kevin Middleton, PhD

Pathology and Anatomical Sciences

Director of Graduate Studies
Associate Professor



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.

Academic Information

Director of Graduate Studies
Associate Professor
P. 573-882-6687

Research Interests

  • Effects of exercise and genetic background on skeletal form, physiology, and biomechanics using high activity mice as a model organism
  • Evolution of the avian head: integrating cranial biomechanics, muscle modeling, and brain evolution to understand avian cranial kinesis
  • Development of Bayesian and machine learning models for the prediction of craniofacial growth and form in humans
  • Evolution of wing-propelled diving in birds: exploring patterns of evolution in bone microstructure, feather biomechanics, functional morphology, and sensory systems

Education & Training

Post-Graduate School

Ecology and Evolutionary Biology, Brown University


  • 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.
  • 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.
  • 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.
  • 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.
  • 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.