Andrew J. Spence, PhD is an Associate Professor in the Department of Bioengineering at Temple University. Andrew is an applied physicist by training who leads a research group in locomotor neuromechanics and spinal cord injury. As a group, the Spence Lab is focused on how the nervous and mechanical systems work together to produce movement, taking an integrative approach that combines experimental work with mathematical modeling, instrumentation, and some robotics. Andrew did his undergraduate work in physics at UC Berkeley, before doing a PhD in neuroscience and biomedical microdevices at Cornell University with Michael Isaacson and Ron Hoy. He returned to Berkeley for a postdoc, and worked with Bob Full on the control of many-legged locomotion, and Eileen Hebets on the neurophysiology of antennas. Before coming to Temple University, he was an RCUK Fellowship awardee and subsequently faculty member in the Structure and Motion Laboratory at the Royal Veterinary College, London, working with Alan Wilson before becoming an independent researcher. Currently his group is focused on applying new neurogenetic techniques (chemogenetics in the form of DREADDs; optogenetics) to 1) better treat spinal cord injuries, 2) dissect the control of fast legged locomotion, and 3) understand how constraints (stability, energetics) have shaped quadrupedal gait control.

Research Interests

  • Movement Science including Control of Locomotion, Gait Analysis, Spinal Cord Injury, Neuroprosthetics, Neuromechanics, Biomechanics, Use of Molecular Genetic Tools (Optogenetics and Chemogenetics/DREADDs
  • Robotics
  • Motor Control
  • Animal Locomotion

Courses Taught




BIOE 4431

Capstone Elective: Neuroengineering


BIOE 4441

Capstone Elective: Biomechanics


BIOE 5431



BIOE 5441



Selected Publications

  • Moukarzel, G., Rauscher, B.C., Patil, C.A., & Spence, A.J. (2022). Infrared videography of a subcutaneous knee tattoo as a simple and inexpensive method to overcome skin motion artifact in rodent kinematics. Cold Spring Harbor Laboratory. doi: 10.1101/2022.08.22.504813

  • Spence, A.J., Wilshin, S.D., & Byrnes, G. (2022). The economy of terrestrial locomotion. Curr Biol, 32(12), pp. R676-R680. England. doi: 10.1016/j.cub.2022.04.063

  • Vahedipour, A., Short, M.R., Timnak, A., Maghsoudi, O.H., Hallowell, T., Gerstenhaber, J., Cappellari, O., Lemay, M., & Spence, A.J. (2021). A versatile system for neuromuscular stimulation and recording in the mouse model using a lightweight magnetically coupled headmount. J Neurosci Methods, 362, p. 109319. Netherlands. doi: 10.1016/j.jneumeth.2021.109319

  • Moukarzel, G., Lemay, M.A., & Spence, A.J. (2021). A MATLAB application for automated H-Reflex measurements and analyses. Biomed Signal Process Control, 66. England. doi: 10.1016/j.bspc.2021.102448

  • Wilshin, S., Reeve, M.A., & Spence, A.J. (2021). Dog galloping on rough terrain exhibits similar limb co-ordination patterns and gait variability to that on flat terrain. Bioinspir Biomim, 16(1), p. 015001. England. doi: 10.1088/1748-3190/abb17a

  • Eisdorfer, J.T., Phelan, M.A., Keefe, K.M., Rollins, M.M., Campion, T.J., Rauscher, K.M., Sobotka-Briner, H., Senior, M., Gordon, G., Smith, G.M., & Spence, A.J. (2021). Addition of angled rungs to the horizontal ladder walking task for more sensitive probing of sensorimotor changes. PLoS One, 16(2), p. e0246298. United States. doi: 10.1371/journal.pone.0246298

  • Tang, Y., Chi, Y., Sun, J., Huang, T., Maghsoudi, O.H., Spence, A., Zhao, J., Su, H., & Yin, J. (2020). Leveraging elastic instabilities for amplified performance: Spine-inspired high-speed and high-force soft robots. Sci Adv, 6(19), p. eaaz6912. United States. doi: 10.1126/sciadv.aaz6912

  • Maghsoudi, O.H. & Spence, A. (2020). Treadmill Training Effect on Kinematics: An Aging Study in Rats. Journal of Medical and Biological Engineering, 40(1), pp. 11-23. doi: 10.1007/s40846-019-00490-x

  • Eisdorfer, J.T., Smit, R.D., Keefe, K.M., Lemay, M.A., Smith, G.M., & Spence, A.J. (2020). Epidural Electrical Stimulation: A Review of Plasticity Mechanisms That Are Hypothesized to Underlie Enhanced Recovery From Spinal Cord Injury With Stimulation. Front Mol Neurosci, 13, p. 163. Switzerland. doi: 10.3389/fnmol.2020.00163

  • Maghsoudi, O.H., Vahedipour, A., & Spence, A. (2019). A novel method for robust markerless tracking of rodent paws in 3D. Eurasip Journal on Image and Video Processing, 2019(1). doi: 10.1186/s13640-019-0477-9

  • Maghsoudi, O.H., Vahedipour, A., & Spence, A. (2019). Three-dimensional-based landmark tracker employing a superpixels method for neuroscience, biomechanics, and biology studies. International Journal of Imaging Systems and Technology, 29(4), pp. 419-430. doi: 10.1002/ima.22317

  • Davies, Z.T.S., Spence, A.J., & Wilson, A.M. (2019). Ground reaction forces of overground galloping in ridden Thoroughbred racehorses. J Exp Biol, 222(Pt 16). England. doi: 10.1242/jeb.204107

  • Maghsoudi, O.H., Vahedipour, A., Hallowell, T., & Spence, A. (2019). Open-source Python software for analysis of 3D kinematics from quadrupedal animals. Biomedical Signal Processing and Control, 51, pp. 364-373. doi: 10.1016/j.bspc.2019.02.024

  • Davies, Z.T.S., Spence, A.J., & Wilson, A.M. (2019). External mechanical work in the galloping racehorse. Biol Lett, 15(2), p. 20180709. England. doi: 10.1098/rsbl.2018.0709

  • Spence, A. (2018). Not just great TV — broad insights from extreme animal performance Feats of Strength: How Evolution Shapes Animal Athletic Abilities Simon Lailvaux (Yale University Press, New Haven and London; 2018) ISBN: 978-0-300-22259-3. Current Biology, 28(20), pp. r1176-r1177. doi: 10.1016/j.cub.2018.09.009

  • Wilshin, S., Shamble, P.S., Hovey, K.J., Harris, R., Spence, A.J., & Hsieh, S.T. (2018). Limping following limb loss increases locomotor stability. J Exp Biol, 221(Pt 18). England. doi: 10.1242/jeb.174268

  • Vahedipour, A., Maghsoudi, O.H., Wilshin, S., Shamble, P., Robertson, B., & Spence, A. (2018). Uncovering the structure of the mouse gait controller: Mice respond to substrate perturbations with adaptations in gait on a continuum between trot and bound. J Biomech, 78, pp. 77-86. United States. doi: 10.1016/j.jbiomech.2018.07.020

  • Maghsoudi, O.H., Vahedipour, A., Gerstenhaber, J., George, S.P., Hallowell, T., Robertson, B., Short, M., & Spence, A. (2018). Matlab software for impedance spectroscopy designed for neuroscience applications. J Neurosci Methods, 307, pp. 70-83. Netherlands. doi: 10.1016/j.jneumeth.2018.06.020

  • Maghsoudi, O.H., Vahedipour, A., Robertson, B., & Spence, A. (2018). Application of Superpixels to Segment Several Landmarks in Running Rodents. Pattern Recognition and Image Analysis, 28(3), pp. 468-482. doi: 10.1134/S1054661818030082

  • Maghsoudi, O.H., Tabrizi, A.V., Robertson, B., & Spence, A. (2018). Superpixels based marker tracking vs. hue thresholding in rodent biomechanics application. Conference Record of 51st Asilomar Conference on Signals, Systems and Computers, ACSSC 2017, 2017-October, pp. 209-213. doi: 10.1109/ACSSC.2017.8335168

  • Wilshin, S., Haynes, G.C., Porteous, J., Koditschek, D., Revzen, S., & Spence, A.J. (2017). Morphology and the gradient of a symmetric potential predict gait transitions of dogs. Biol Cybern, 111(3-4), pp. 269-277. Germany. doi: 10.1007/s00422-017-0721-2

  • Maghsoudi, O.H., Tabrizi, A.V., Robertson, B., & Spence, A. (2017). 3D modeling of running rodents based on direct linear transform. 2017 IEEE Signal Processing in Medicine and Biology Symposium, SPMB 2017 - Proceedings, 2018-January, pp. 1-4. doi: 10.1109/SPMB.2017.8257039

  • Maghsoudi, O.H., Hallowell, T., Tabrizi, A.V., George, S.P., Robertson, B., Short, M., Gerstenhaber, J., & Spence, A. (2017). Impedance characterization of bipolar implantable nerve cuffs for neuroscience applications. 2017 IEEE Signal Processing in Medicine and Biology Symposium, SPMB 2017 - Proceedings, 2018-January, pp. 1-3. doi: 10.1109/SPMB.2017.8257040

  • Wilshin, S., Reeve, M.A., Haynes, G.C., Revzen, S., Koditschek, D.E., & Spence, A.J. (2017). Longitudinal quasi-static stability predicts changes in dog gait on rough terrain. J Exp Biol, 220(Pt 10), pp. 1864-1874. England. doi: 10.1242/jeb.149112

  • Hsieh, S.T., Shamble, P., Wilshin, S., Hovey, K., & Spence, A.J. (2017). Spiders "limp" to achieve a more stable gait. INTEGRATIVE and COMPARATIVE BIOLOGY, 57, pp. E297-E297. Retrieved from

  • Maghsoudi, O.H., Tabrizi, A.V., Robertson, B., Shamble, P., & Spence, A. (2017). A Rodent Paw Tracker Using Support Vector Machine. 2016 IEEE Signal Processing in Medicine and Biology Symposium, SPMB 2016 - Proceedings. doi: 10.1109/SPMB.2016.7846866

  • Spence, A. & IEEE (2017). Gait, posture, pogo-sticks and newfangled neurogenetics: How do legged animals control their locomotion? 2017 IEEE SIGNAL PROCESSING in MEDICINE and BIOLOGY SYMPOSIUM (SPMB). Retrieved from

  • Charles, J.P., Cappellari, O., Spence, A.J., Wells, D.J., & Hutchinson, J.R. (2016). Muscle moment arms and sensitivity analysis of a mouse hindlimb musculoskeletal model. J Anat, 229(4), pp. 514-535. England. doi: 10.1111/joa.12461

  • Maghsoudi, O.H., Tabrizi, A.V., Robertson, B., Shamble, P., & Spence, A. (2016). A novel automatic method to track the body and paws of running mice in high speed video. 2015 IEEE Signal Processing in Medicine and Biology Symposium - Proceedings. doi: 10.1109/SPMB.2015.7405456

  • Charles, J.P., Cappellari, O., Spence, A.J., Hutchinson, J.R., & Wells, D.J. (2016). Musculoskeletal Geometry, Muscle Architecture and Functional Specialisations of the Mouse Hindlimb. PLoS One, 11(4), p. e0147669. United States. doi: 10.1371/journal.pone.0147669

  • Charles, J., Cappellari, O., Spence, A., Wells, D., & Hutchinson, J. (2015). Developing, Testing and Optimising a Mouse Hindlimb Musculoskeletal Model. FASEB JOURNAL, 29. Retrieved from

  • Wilshin, S.D., Starr, J., Clark, G.C., Koditschek, D.E., & Spence, A.J. (2015). Using a physical model to investigate dog walking behaviour on rough terrain. INTEGRATIVE and COMPARATIVE BIOLOGY, 55, pp. E200-E200. Retrieved from

  • Mamuneas, D., Spence, A.J., Manica, A., & King, A.J. (2015). Bolder stickleback fish make faster decisions, but they are not less accurate. Behavioral Ecology, 26(1), pp. 91-96. doi: 10.1093/beheco/aru160

  • Reeve, M.A., Wilshin, S., & Spence, A.J. (2014). Dog gait on rough terrain: When does static stability matter? INTEGRATIVE and COMPARATIVE BIOLOGY, 54, pp. E337-E337. Retrieved from

  • Byrnes, G., Spence, A.J., Martino, B., Hilt, M., & Wilson, A.M. (2014). The effects of gap distance and substrate compliance on the biomechanics of jumping in gray squirrels (Sciurus carolinensis). INTEGRATIVE and COMPARATIVE BIOLOGY, 54, pp. E247-E247. Retrieved from

  • Spence, A.J., Nicholson-Thomas, G., & Lampe, R. (2013). Closing the loop in legged neuromechanics: an open-source computer vision controlled treadmill. J Neurosci Methods, 215(2), pp. 164-169. Netherlands. doi: 10.1016/j.jneumeth.2013.03.009

  • Spence, A.J., Thurman, A.S., Maher, M.J., & Wilson, A.M. (2012). Speed, pacing strategy and aerodynamic drafting in Thoroughbred horse racing. Biol Lett, 8(4), pp. 678-681. England. doi: 10.1098/rsbl.2011.1120

  • Self, Z.T., Spence, A.J., & Wilson, A.M. (2012). Speed and incline during thoroughbred horse racing: racehorse speed supports a metabolic power constraint to incline running but not to decline running. J Appl Physiol (1985), 113(4), pp. 602-607. United States. doi: 10.1152/japplphysiol.00560.2011

  • Spence, A.J. & Hutchinson, J.R. (2012). A growing size synthesis. Curr Biol, 22(9), pp. R309-R314. England. doi: 10.1016/j.cub.2012.03.017

  • Wilshin, S., Haynes, G.C., Reeve, M., Revzen, S., & Spence, A.J. (2012). How is dog gait affected by natural rough terrain? INTEGRATIVE and COMPARATIVE BIOLOGY, 52, pp. E193-E193. Retrieved from

  • Liedtke, A.M., Moore, S., Witte, T., & Spence, A.J. (2012). How do animals with limited distal limb musculature use sensory feedback during locomotion? INTEGRATIVE and COMPARATIVE BIOLOGY, 52, pp. E106-E106. Retrieved from

  • Self, Z.T., Spence, A.J., & Wilson, A.M. (2012). Jump racing: do horses slow down due to a force limit? INTEGRATIVE and COMPARATIVE BIOLOGY, 52, pp. E157-E157. Retrieved from

  • Wilshin, S.D., Haynes, G.C., Porteous, J., & Spence, A.J. (2012). Describing gait transitions and the role of symmetry in control. INTEGRATIVE and COMPARATIVE BIOLOGY, 52, pp. E194-E194. Retrieved from

  • Byrnes, G. & Spence, A.J. (2011). Ecological and biomechanical insights into the evolution of gliding in mammals. Integr Comp Biol, 51(6), pp. 991-1001. England. doi: 10.1093/icb/icr069

  • Byrnes, G., Libby, T., Lim, N.T., & Spence, A.J. (2011). Gliding saves time but not energy in Malayan colugos. J Exp Biol, 214(Pt 16), pp. 2690-2696. England. doi: 10.1242/jeb.052993

  • Sponberg, S., Spence, A.J., Mullens, C.H., & Full, R.J. (2011). A single muscle's multifunctional control potential of body dynamics for postural control and running. Philos Trans R Soc Lond B Biol Sci, 366(1570), pp. 1592-1605. England. doi: 10.1098/rstb.2010.0367

  • Byrnes, G., Lim, N., Yeong, C., & Spence, A.J. (2011). Sex differences in the locomotor ecology of a gliding mammal, the Malayan colugo (Galeopterus variegatus). Journal of Mammalogy, 92(2), pp. 444-451. doi: 10.1644/10-MAMM-A-048.1

  • Byrnes, G. & Spence, A.J. (2011). Ecological and biomechanical insights into the origins of gliding in mammals. INTEGRATIVE and COMPARATIVE BIOLOGY, 51, pp. E18-E18. Retrieved from

  • Parsons, K.J., Spence, A.J., Morgan, R., Thompson, J.A., & Wilson, A.M. (2011). High speed field kinematics of foot contact in elite galloping horses in training. Equine Vet J, 43(2), pp. 216-222. United States. doi: 10.1111/j.2042-3306.2010.00149.x

  • Wilshin, S., Kelleher, C.N., Byrnes, G., Seipel, J., & Spence, A.J. (2011). Dogs on springs: do trotting dogs adjust their virtual leg stiffness on compliant surfaces? INTEGRATIVE and COMPARATIVE BIOLOGY, 51, pp. E152-E152. Retrieved from

  • Willemart, R.H., Santer, R.D., Spence, A.J., & Hebets, E.A. (2011). A sticky situation: Solifugids (Arachnida, Solifugae) use adhesive organs on their pedipalps for prey capture. Journal of Ethology, 29(1), pp. 177-180. doi: 10.1007/s10164-010-0222-4

  • Spence, A.J., Revzen, S., Seipel, J., Mullens, C., & Full, R.J. (2010). Insects running on elastic surfaces. J Exp Biol, 213(11), pp. 1907-1920. England. doi: 10.1242/jeb.042515

  • Pfau, T., Spence, A., Starke, S., Ferrari, M., & Wilson, A. (2009). Modern riding style improves horse racing times. Science, 325(5938), p. 289. United States. doi: 10.1126/science.1174605

  • Spence, A.J., Seipel, J., Revzen, S., Mullens, C., Yeats, K., & Full, R.J. (2009). Insects running on elastic surfaces: The role of feedforward control. COMPARATIVE BIOCHEMISTRY and PHYSIOLOGY A-MOLECULAR & INTEGRATIVE PHYSIOLOGY, 153A(2), pp. S137-S137. doi: 10.1016/j.cbpa.2009.04.249

  • Snarey, R.A., Spence, A., & Wilson, A. (2009). The kinematics and kinetics of grey squirrel jumping. COMPARATIVE BIOCHEMISTRY and PHYSIOLOGY A-MOLECULAR & INTEGRATIVE PHYSIOLOGY, 153A(2), pp. S130-S130. doi: 10.1016/j.cbpa.2009.04.228

  • Spence, A.J., Thurman, A., Maher, M., & Wilson, A.M. (2009). Speed, strategy, drag and drafting in thoroughbred horse racing. COMPARATIVE BIOCHEMISTRY and PHYSIOLOGY A-MOLECULAR & INTEGRATIVE PHYSIOLOGY, 153A(2), pp. S127-S127. doi: 10.1016/j.cbpa.2009.04.216

  • Self, Z.T., Spence, A.J., Bowtell, M., & Wilson, A.M. (2009). Does running downhill affect maximum speed? COMPARATIVE BIOCHEMISTRY and PHYSIOLOGY A-MOLECULAR & INTEGRATIVE PHYSIOLOGY, 153A(2), pp. S132-S132. doi: 10.1016/j.cbpa.2009.04.233

  • Byrnes, G., Lim, N., & Spence, A.J. (2009). Integrating locomotor mechanics and transport costs in a free-ranging gliding mammal. INTEGRATIVE and COMPARATIVE BIOLOGY, 49, pp. E24-E24. Retrieved from

  • Spence, A.J. (2009). Scaling in biology. Curr Biol, 19(2), pp. R57-R61. England. doi: 10.1016/j.cub.2008.10.042

  • Spence, A.J., Tan, H., & Wilson, A. (2008). Accuracy of the TurfTrax Racing Data System for determination of equine speed and position. Equine Vet J, 40(7), pp. 680-683. United States. doi: 10.2746/042516408x330338

  • Byrnes, G., Lim, N.T., & Spence, A.J. (2008). Take-off and landing kinetics of a free-ranging gliding mammal, the Malayan colugo (Galeopterus variegatus). Proc Biol Sci, 275(1638), pp. 1007-1013. England. doi: 10.1098/rspb.2007.1684

  • Spence, A.J., Neeves, K.B., Murphy, D., Sponberg, S., Land, B.R., Hoy, R.R., & Isaacson, M.S. (2007). Flexible multielectrodes can resolve multiple muscles in an insect appendage. J Neurosci Methods, 159(1), pp. 116-124. Netherlands. doi: 10.1016/j.jneumeth.2006.07.002

  • Spence, A.J. & Hebets, E.A. (2006). Anatomy and physiology of giant neurons in the antenniform leg of the amblypygid Phrynus marginemaculatus. Journal of Arachnology, 34(3), pp. 566-577. doi: 10.1636/S05-53.1

  • Spence, A.J., Revzen, S., Yeates, K., Mullens, C., & Full, R.J. (2006). Insects running on compliant surfaces. INTEGRATIVE and COMPARATIVE BIOLOGY, 46, pp. E134-E134. Retrieved from

  • Byrnes, G.T. & Spence, A.J. (2006). Locomotor biomechanics of a free-ranging gliding mammal (Cynocephalus variegatus). INTEGRATIVE and COMPARATIVE BIOLOGY, 46, pp. E20-E20. Retrieved from

  • Revzen, S., Bishop-Moser, J., Spence, A., & Full, R.J. (2006). Testing control models in rapid running insects using lateral ground translation. INTEGRATIVE and COMPARATIVE BIOLOGY, 46, pp. E116-E116. Retrieved from

  • Spataro, L., Dilgen, J., Retterer, S., Spence, A.J., Isaacson, M., Turner, J.N., & Shain, W. (2005). Dexamethasone treatment reduces astroglia responses to inserted neuroprosthetic devices in rat neocortex. Exp Neurol, 194(2), pp. 289-300. United States. doi: 10.1016/j.expneurol.2004.08.037

  • Retterer, S.T., Smith, K.L., Bjornsson, C.S., Neeves, K.B., Spence, A.J.H., Turner, J.N., Shain, W., & Isaacson, M.S. (2004). Model neural prostheses with integrated microfluidics: a potential intervention strategy for controlling reactive cell and tissue responses. IEEE Trans Biomed Eng, 51(11), pp. 2063-2073. United States. doi: 10.1109/TBME.2004.834288

  • Russo, A.P., Retterer, S.T., Spence, A.J., Isaacson, M.S., Lepak, L.A., Spencer, M.G., Martin, D.L., MacColl, R., & Turner, J.N. (2004). Direct casting of polymer membranes into microfluidic devices. Separation Science and Technology, 39(11), pp. 2515-2530. doi: 10.1081/SS-200026706

  • James, C.D., Spence, A.J.H., Dowell-Mesfin, N.M., Hussain, R.J., Smith, K.L., Craighead, H.G., Isaacson, M.S., Shain, W., & Turner, J.N. (2004). Extracellular recordings from patterned neuronal networks using planar microelectrode arrays. IEEE Trans Biomed Eng, 51(9), pp. 1640-1648. United States. doi: 10.1109/TBME.2004.827252

  • Szarowski, D.H., Andersen, M.D., Retterer, S., Spence, A.J., Isaacson, M., Craighead, H.G., Turner, J.N., & Shain, W. (2003). Brain responses to micro-machined silicon devices. Brain Res, 983(1-2), pp. 23-35. Netherlands. doi: 10.1016/s0006-8993(03)03023-3

  • Surlykke, A., Yack, J.E., Spence, A.J., & Hasenfuss, I. (2003). Hearing in hooktip moths (Drepanidae: Lepidoptera). J Exp Biol, 206(Pt 15), pp. 2653-2663. England. doi: 10.1242/jeb.00469

  • Spence, A.J., Hoy, R.R., & Isaacson, M.S. (2003). A micromachined silicon multielectrode for multiunit recording. J Neurosci Methods, 126(2), pp. 119-126. Netherlands. doi: 10.1016/s0165-0270(03)00075-x

  • Turner, J.N., Szarowski, D.H., Shain, W., Buttle, K., Tivol, W.F., Bagle, H., Spence, A.J., Retterer, S., Lapek, L., Richards, T., Isaacson, M., & Spencer, M. (2002). "On-chip" NanoFabricated collagen membranes observed by high-voltage electron microscopy. Microscopy and Microanalysis, 8(SUPPL. 2), pp. 1118-1119. doi: 10.1017/s1431927602103667

  • Krivanek, O.L., Dellby, N., Spence, A.J., & Brown, L.M. (1998). Spherical aberration correction in dedicated STEM. ELECTRON MICROSCOPY 1998, VOL 1, pp. 55-56. Retrieved from

  • Krivanek, O.L., Dellby, N., Spence, A.J., Camps, R.A., & Brown, L.M. (1997). Aberration correction in the STEM. ELECTRON MICROSCOPY and ANALYSIS 1997(153), pp. 35-40. Retrieved from