Department: Comparative Biomedical Sciences

Campus: Hawkshead

Research Groups: Brain Health and Behaviour, Musculoskeletal Biology

Research Centres: RVC Quantitative Biology Resource, Structure & Motion Laboratory

Dr Monica Daley is a Senior Lecturer in Locomotor Biomechanics. Her research team investigate how animals achieve agile legged locomotion in complex environments.

Monica earned an HBSc in Biology with a Chemistry minor at the University of Utah, where she was inspired to pursue an academic career through her research on human locomotor-ventilatory integration with Dennis Bramble and David Carrier. Monica then spent a year as a research technician at the University of Utah, investigating motor control of singing in zebra finches in the lab of Franz Goller. These experiences initiated a long-standing fascination with the interplay of mechanics and neural control.

Monica went on to Harvard University, where she earned her MA and PhD in Organismic and Evolutionary Biology. Her research on muscle-tendon dynamics and biomechanics of avian bipedal locomotion was supported by a prestigious Predoctoral Fellowship award from the Howard Hughes Medical Institute and supervised by Andrew Biewener at the Concord Field Station of Harvard University (CFS Website).

After completing her PhD, Monica was awarded a Research Fellowship by the U.S. National Science Foundation to develop models of the dynamics and control of bipedal locomotion, working with Dan Ferris in the Human Neuromechanics Lab at University of Michigan, in collaboration with Auke Ijspeert in the Biologically Inspired Robotics Group at the Swiss Federal Institute of Technology in Lausanne.

Monica is a faculty member of the Structure and Motion Lab, where she leads research in Comparative Neuromechanics— a field that seeks to understand the interplay of morphology, mechanics and sensorimotor control that influences how animals move through their environment.

Monica serves as Associate Editor for Royal Society Open Science and on the Editorial Board for Journal of Experimental Biology and Physiological and Biochemical Zoology.  Daley served as a Guest Editor for Frontiers in Robotics and AI, for a theme issue on Advances in Mechatronics and Biomechanics towards Efficient Robot Actuation.  She serves on the Scientific Board of the conferences Adaptive Motion in Animals and Machines and Dynamic Walking and the Steering Committee for the London Centre for Ecology and Evolution. Monica is a Fellow of The Higher Education Academy and a member of several professional societies including the Society for Integrative and Comparative Biology, Society for Experimental Biology and the American Physiological Society. 

Daley has given 22 keynote and invited talks over the past 5 years. 


Locomotion is a key behaviour for most animals, including humans. To survive and reproduce, animals must move through complex environments to obtain food, escape predators, and find mates. Animal locomotion often involves impressive athletic feats such as long distance migration, high speeds, and rapid manoeuvring. Animals accomplish a wide range of behaviours with a single physiological system and yet still often out-perform human-engineered machines.

Dr Daley an integrative physiologist, broadly interested in the sensory, mechanical, neuromuscular and energetic factors that influence animal locomotion. Daley’s research team investigate how animals achieve robustly stable, economic and agile locomotion in complex terrain. This research is at the interface of muscle physiology, energetics, neural control, mechanics and bio-inspired robotics.

We use an integrative approach that combines a range of experimental techniques in biomechanics and physiology with computer simulations and collaborative bio-inspired design and control of legged robots. We work with engineers to translate biological principles into technological applications.

We often use ground birds as a model system because birds are diverse bipedal athletes that span a large size range from tiny 35 gram painted quail to ostriches over 100 kg. Yet, ground birds use striding bipedal gaits that are mechanically similar to human walking and running. By studying ground birds, we can reveal how muscle-tendon architecture, leg morphology, body size and terrain environment influence bipedal locomotion.

This research provides insight into natural animal behaviour, evolution of leg morphology, and injury mechanisms in domestic and wild species. Such insights can inform health and welfare issues including mobility throughout the lifespan, gait pathology, fall prevention and rehabilitation.

Alongside research in comparative animal locomotion, Daley maintains interests in human locomotion, including biomechanics, energetics, locomotor-ventilatory interactions and rehabilitation approaches informed by neuromechanical principles.

For more information about Daley’s recent work, see: 

Structure & Motion Lab

Activity and behavior monitoring in pets:  New industry partnership project between RVC and Felcana pet technology.

Featured article in Journal of Experimental Biology: Ostriches specialized for economic running

Secretary Birds can kick a snake to death in the blink of an eye

RVC Projects: Don’t Break a Leg!

Running birds prioritise safety on uneven terrain

Teaching robots to run like birds

Does your breathing pattern matter?

'ATRIAS@RVC' blog on the robotics experiments with our engineering collaborators from Dr Jonathan Hurst's Dynamic Robotics Laboratory at Oregon State University (funded by Human Frontier Science Program).

'Towards the Chicken of the Future' blog about collaborative work on broiler chicken breathing and locomotion, led by Professor John Hutchinson and Dr Jonathan Codd

For a continuously updated list of publications,  see Dr Daley' s Google Scholar Profile

Daley (2018) Understanding the agility of running birds: Sensorimotor and mechanical factors in avian bipedal locomotion. Integrative and Comparative Biology.

Neal, Barton, Birn-Jeffrey, Daley, Morrissey (2018) The effects & mechanisms of increasing running step rate: A feasibility study in a mixed-sex group of runners with patellofemoral pain. Physical Therapy in Sport. May 2018

Daley and Birn-Jeffery (2018) Scaling of avian bipedal locomotion reveals independent effects of body mass and leg posture on gait. J Exp Bio. 221(10):jeb152538. Altmetrics in top 5%, ranked most read in journal June-October 2018.

Urbina-Melendez, Jalaledini, Daley, Valero-Cuevas (2018). A physical model suggests that hip-localized balance sense in birds improves state estimation for perching: implications for bipedal robots. Frontiers in Robotics and AI, 5, p.38.

Daley, Channon, Nolan, Hall (2016) Preferred gait and walk-run transition speeds in ostriches measured using GPS-IMU sensors. J Exp Bio, 219: 3301-3308. Featured article in Inside JEB, see:

Portugal, Murn, Sparkes, Daley (2016) The fast and forceful kicking strike of the secretary bird. Current Biology, Volume 26, Issue 2, R58-R59. Featured in BBC Science and Environment:

Daley (2016) Non-steady Locomotion, in Understanding Mammalian Locomotion: Concepts and Applications (ed J. E. A. Bertram), John Wiley & Sons, Inc, Hoboken, NJ.

Gordon, Rankin, Daley (2015) How do treadmill speed and terrain visibility influence neuromuscular control of guinea fowl locomotion? J Exp Bio, 218(19), 3010-3022.

Hubicki, Jones, Daley, Hurst (2015). Do limit cycles matter in the long run? Stable orbits and sliding-mass dynamics emerge in task-optimal locomotion. IEEE International Conference on Robotics and Automation (ICRA), 5113-5120.

Birn-Jeffery A.V., Hubicki C., Blum Y., Renjewski D., Hurst J.W. and Daley M.A. (2014) Don't break a leg: Running birds from quail to ostrich prioritize leg safety and economy in uneven terrain. Journal of Experimental Biology. J Exp Biol 217, 3786-3796. Featured article in Inside JEB, see:

Blum Y., Vejdani H.R., Birn-Jeffery A. V., Hubicki C. , Hurst J.W. and Daley M.A. (2014) Swing-Leg Trajectory of Running Guinea Fowl Suggests Task-Level Priority of Force Regulation Rather than Disturbance Rejection. PLoS ONE 9(6): e100399.

Van Why J., Hubiki C., Jones M., Daley M. and Hurst J. (2014) Running into a Trap: Numerical Design of Task-Optimal Preflex Behaviors for Delayed Disturbance Responses. IEEE International Conference on Intelligent Robots and Systems (IROS) (DOI: 10.1109/IROS.2014.6942908), Sept. 2014.

Vejdani H.R., Blum Y., Daley M.A., and Hurst J.W. (2013) Bio-inspired swing leg control for spring-mass robots running on ground with unexpected height disturbance, Bioinspiration & Biomimetics (DOI:10.1088/1748-3182/8/4/046006)

Daley M.A., Bramble D.M., and Carrier D.R. (2013). Impact loading and locomotor-respiratory coordination significantly influence breathing dynamics in running humans. PLOS ONE. August 2013 8(8) e70752  DOI:10.1371/journal.pone.0070752

Voloshina A.S., Kuo A.D., Daley M.A. Ferris D.P. (2013) Biomechanics and energetics of walking on uneven terrain. Journal of Experimental Biology. August, 2013 DOI:10.1242/jeb.081711 

Paxton H., Daley M., Corr S., Hutchinson J. (2013) The gait dynamics of the modern broiler chicken: A cautionary tale of selective breeding. Journal of Experimental Biology. DOI:10.1242/jeb.080309

Ross C.F., Blob R.W., Carrier D.R., Daley M.A., Deban S.M., Demes B., Gripper J., Iriarte-Diaz J., Kilbourne B.M., Landberg T., Polk J.D., Schilling N., Vanhooydonck B. (2012) Evolution of locomotor rhythmicity in tetrapods. Evolution. DOI:10.1111/evo.12015

BIRN-JEFFERY, A.V. and DALEY, M.A. (2012). Birds achieve high robustness in uneven terrain through active control of landing conditions. J Exp Bio 215, 2117-2127.

BLUM, Y.N., BIRN-JEFFERY, A. DALEY, M.A. and SEYFARTH, A. (2011). Does A Crouched Leg Posture Enhance Running Stability and Robustness? Journal of Theoretical Biology. 281, 97-106.

DALEY, M.A., and BIEWENER, A.A. (2011). Leg muscles that mediate stability: Mechanics and control of two distal extensor muscles during obstacle negotiation in the guinea fowl. Phil Trans R Soc B. 366, 1580-1591. DOI: 10.1098/rstb.2010.0338.

DALEY, M. A. & USHERWOOD, J. R. (2010) Two explanations for the compliant running paradox: reduced work of bouncing viscera and increased stability in uneven terrain. Biol Lett. 6 (3) p. 418-21 doi:10.1098/rsbl.2010.0175. Open access: see the Full Paper and Online Supplementary Material.

DALEY, M. A., VOLOSHINA, A. & BIEWENER, A. A. (2009) The role of intrinsic muscle mechanics in the neuromuscular control of stable running in the guinea fowl. J Physiol 587, 2693-2707. PubMed ID 19359369

DALEY, M. A. (2008) Biomechanics: running over uneven terrain is a no-brainer. Curr Biol 18, R1064-1066. PubMed ID 19036337

PELC, E. H., DALEY, M. A. & FERRIS, D. P. (2008) Resonant hopping of a robot controlled by an artificial neural oscillator. Bioinspiration and Biomimetics. 3, 26001. PubMed ID 18369282

FERRIS, D. P., SAWICKI, G. S. & DALEY, M. A. (2007) A physiologist's perspective on robotic exoskeletons for human locomotion. International Journal of Humanoid Robotics 4, 507-528. PubMed ID 18185840

DALEY, M. A., FELIX, G. & BIEWENER, A. A. (2007) Running stability is enhanced by a proximo-distal gradient in joint neuromechanical control. Journal of Experimental Biology 210, 383-394. PubMed ID 17234607

BIEWENER, A. A. & DALEY, M. A. (2007) Unsteady locomotion: integrating muscle function with whole body dynamics and neuromuscular control. Journal of Experimental Biology 210, 2949-2960. PubMed ID 17704070

DALEY, M. A., USHERWOOD, J. R., FELIX, G. & BIEWENER, A. A. (2006) Running over rough terrain: guinea fowl maintain dynamic stability despite a large unexpected change in substrate height. Journal of Experimental Biology 209, 171-187. PubMed ID 16354788

DALEY, M. A. & BIEWENER, A. A. (2006) Running over rough terrain reveals limb control for intrinsic stability. Proceedings of the National Academy of Sciences of the United States of America 103, 15681-15686. PubMed ID 17032779

DALEY, M. A. & BIEWENER, A. A. (2003) Muscle force-length dynamics during level versus incline locomotion: a comparison of in vivo performance of two guinea fowl ankle extensors. Journal of Experimental Biology 206, 2941-2958.PubMed ID 12878663

GOLLER, F. & DALEY, M. A. (2001) Novel motor gestures for phonation during inspiration enhance the acoustic complexity of birdsong. Proceedings of the Royal Society of London Series B-Biological Sciences 268, 2301-2305. PubMed ID 11703869


Dr Daley is passionate about teaching, mentoring and outreach, and in particular, encouraging equal opportunities and diverse participation in the sciences.  Daley teaches comparative physiology and musculoskeletal biology topics, as well as research skills including study design and methods, data analysis, statistics, Matlab programming, scientific writing and presentation.

Daley is the module leader for The Moving Animal, part of the Essential Biomedical Sciences taught to students in the Gateway Programme and the 1st year of the BSc degrees. She also leads the Biomechanics and Neural Control research theme for 2nd year BSc research projects and mentors 3rd year undergraduate research projects.  Daley also teaches locomotion topics on the BVetMed course and in an advanced module in the 3rd year of the BSc Bioveterinary Sciences, Comparative Animal Locomotion.

Daley is also actively involved in research training of graduate students. She serves on the management committee of the London Interdisciplinary Doctoral Biosciences Consortium, organises the RVC Postgraduate Research Seminar Series, and leads training workshops for PhD students in the Structure & Motion Lab.

Recent outreach activities:

Consultation and demonstration of avian anatomy and biomechanics for a BBC 4 documentary ‘Can Science Make Me Perfect’, with Professor Alice Roberts.

Interviewed on cross-disciplinary collaborative research, Robohub podcast: Learning about legged locomotion birds

Coding workshop for Researc/hers code:  Modelling Animal Locomotion: A Biologist's Journey. 


Current and upcoming external activities: 

Dr Daley and Dr Birn-Jeffery (QMUL) are co-organising a symposium on Neuromechanics of Legged Locomotion at the next annual meeting for the Society of Experimental Biology,  in Seville Spain, 2-5th July 2019. 

Neuromechanics and Neurophysics: Sensory and mechanical factors underlying stable and agile control of legged locomotion. 

If you are interested in contributing, please submit an abstract through the SEB meeting pages (Deadline 22nd March 2019). 


Dr Daley and Dr McGowan (U of Idaho) are co-organising a symposium on Comparative biomechanics at the upcoming Congress of the International Society of Biomechanics at Calgary from 31 July - 4 Aug 2019. 


Dr Daley is on the Program Committee for the upcoming conference on Adaptive Motion in Animals and Machines,  to be held at EPFL in Lausanne Switzerland, 19th-23rd August 2019. 


If you interested in participating in the above meetings and have any questions, please get in touch. 

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