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The energetic cost of human walking as a function of uneven terrain amplitude
Open accessPublished version of a preprint
Abstract
Humans expend more energy walking on uneven terrain, but the amount varies across terrains. Few experimental characterizations exist, each describing terrain qualitatively without any relation to others or to flat ground. This precludes mechanistic explanation of the energy costs. Here, we show that energy costs vary smoothly and approximately quadratically as a function of terrain amplitude. We tested this with healthy adults (N=10) walking on synthetic uneven terrain with random step heights of parametrically controlled maximum amplitude (four conditions 0–0.045 m) and at four walking speeds (0.8–1.4 m s−1). Both net metabolic rate and the rate of positive work increased approximately with amplitude squared and speed cubed (R2=0.74, 0.82, respectively), as predicted by a simple walking model. The model requires work to redirect the body center of mass velocity between successive arcs described by pendulum-like legs, at proportional metabolic cost. Humans performed most of the greater work with terrain amplitude early in the single stance phase, and with speed later in stance during push-off. Work and energy rates changed with approximately linear proportionality, with a ratio or delta efficiency of 49.5% (R2=0.68). The efficiency was high enough to suggest substantial work performed passively by elastic tendon and not only by active muscle. Simple kinematic measures such as mid-swing foot clearance also increased with terrain amplitude (R2=0.65), possibly costing energy as well. Nevertheless, most of the metabolic cost of walking faster or on more uneven terrain can be explained mechanistically by the work performed.
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Citations
Cites
A general and simple method for obtaining R 2 from generalized linear mixed‐effects models
Nakagawa, S., & Schielzeth, H. (2013) · Methods in Ecology and Evolution · doi:10.1111/j.2041-210x.2012.00261.x · Cited by 10,356
Statsmodels: Econometric and Statistical Modeling with Python
Seabold, S., & Perktold, J. (2010) · Proceedings of the Python in Science Conferences · doi:10.25080/majora-92bf1922-011 · Cited by 6,968
Zeni, J. A., Richards, J. G., & Higginson, J. S. (2008) · Gait & Posture · doi:10.1016/j.gaitpost.2007.07.007 · Cited by 1,261
Active control of lateral balance in human walking
Bauby, C. E., & Kuo, A. D. (2000) · Journal of Biomechanics · doi:10.1016/s0021-9290(00)00101-9 · Cited by 851
Self-citationEnergetic aspects of muscle contraction
WOLEDGE, R. C. (1986) · General Pharmacology The Vascular System · doi:10.1016/0306-3623(86)90238-7 · Cited by 753
Energetics of Actively Powered Locomotion Using the Simplest Walking Model
Kuo, A. D. (2001) · Journal of Biomechanical Engineering · doi:10.1115/1.1427703 · Cited by 740
Self-citationEnergetic Consequences of Walking Like an Inverted Pendulum: Step-to-Step Transitions
Kuo, A. D., Donelan, J. M., & Ruina, A. (2005) · Exercise and Sport Sciences Reviews · doi:10.1097/00003677-200504000-00006 · Cited by 675
Self-citationDonelan, J. M., Kram, R., & Kuo, A. D. (2002) · Journal of Experimental Biology · doi:10.1242/jeb.205.23.3717 · Cited by 670
Self-citationMechanical work and efficiency in level walking and running
Cavagna, G. A., & Kaneko, M. (1977) · The Journal of Physiology · doi:10.1113/jphysiol.1977.sp011866 · Cited by 664
Local Dynamic Stability Versus Kinematic Variability of Continuous Overground and Treadmill Walking
Dingwell, J. B., Cusumano, J. P., Cavanagh, P. R., & Sternad, D. (2000) · Journal of Biomechanical Engineering · doi:10.1115/1.1336798 · Cited by 664
Stabilization of Lateral Motion in Passive Dynamic Walking
Kuo, A. D. (1999) · The International Journal of Robotics Research · doi:10.1177/02783649922066655 · Cited by 659
Self-citationDerivation of formulae used to calculate energy expenditure in man.
Brockway, J. M. (1987) · PubMed · Cited by 567
Mechanical and metabolic determinants of the preferred step width in human walking
Maxwell Donelan, J., Kram, R., & Arthur D., K. (2001) · Proceedings of the Royal Society of London. Series B: Biological Sciences · doi:10.1098/rspb.2001.1761 · Cited by 552
Self-citationSimultaneous positive and negative external mechanical work in human walking
Donelan, J. M., Kram, R., & Kuo, A. D. (2002) · Journal of Biomechanics · doi:10.1016/s0021-9290(01)00169-5 · Cited by 530
Self-citationMechanical work in terrestrial locomotion: two basic mechanisms for minimizing energy expenditure
Cavagna, G. A., Heglund, N. C., & Taylor, C. R. (1977) · American Journal of Physiology-Regulatory, Integrative and Comparative Physiology · doi:10.1152/ajpregu.1977.233.5.r243 · Cited by 515
Mechanical and metabolic requirements for active lateral stabilization in human walking
Donelan, J. M., Shipman, D. W., Kram, R., & Kuo, A. D. (2004) · Journal of Biomechanics · doi:10.1016/j.jbiomech.2003.06.002 · Cited by 491
Self-citationA Simple Model of Bipedal Walking Predicts the Preferred Speed–Step Length Relationship
Kuo, A. D. (2001) · Journal of Biomechanical Engineering · doi:10.1115/1.1372322 · Cited by 448
Self-citationFORCES AND ENERGY CHANGES IN THE LEG DURING WALKING
Elftman, H. (1939) · American Journal of Physiology-Legacy Content · doi:10.1152/ajplegacy.1939.125.2.339 · Cited by 338
Gard, S. A., Miff, S. C., & Kuo, A. D. (2004) · Human Movement Science · doi:10.1016/j.humov.2003.11.002 · Cited by 325
Self-citationProceedings of the 19th Python in Science Conference
(2020) · Proceedings of the Python in Science Conferences · doi:10.25080/majora-342d178e-02b · Cited by 285
Positive and negative work performances and their efficiencies in human locomotion
Margaria, R. (1968) · European Journal of Applied Physiology · doi:10.1007/bf00699624 · Cited by 276
The advantages of a rolling foot in human walking
Adamczyk, P. G., Collins, S. H., & Kuo, A. D. (2006) · Journal of Experimental Biology · doi:10.1242/jeb.02455 · Cited by 271
Self-citationWang, Y., & Srinivasan, M. (2014) · Biology Letters · doi:10.1098/rsbl.2014.0405 · Cited by 265
The Effect of Lateral Stabilization on Walking in Young and Old Adults
Dean, J. C., Alexander, N. B., & Kuo, A. D. (2007) · IEEE Transactions on Biomedical Engineering · doi:10.1109/tbme.2007.901031 · Cited by 245
Self-citationMechanics and energetics of swinging the human leg
Doke, J., Donelan, J. M., & Kuo, A. D. (2005) · Journal of Experimental Biology · doi:10.1242/jeb.01408 · Cited by 238
Self-citationEnergy-speed relation and optimal speed during level walking
Ralston, H. J. (1958) · Internationale Zeitschrift für Angewandte Physiologie Einschliesslich Arbeitsphysiologie · doi:10.1007/bf00698754 · Cited by 203
Terrain coefficients for energy cost prediction.
Soule, R. G., & Goldman, R. F. (1972) · Journal of Applied Physiology · doi:10.1152/jappl.1972.32.5.706 · Cited by 202
Biomechanics and energetics of walking on uneven terrain
Voloshina, A. S., Kuo, A. D., Daley, M. A., & Ferris, D. P. (2013) · Journal of Experimental Biology · doi:10.1242/jeb.081711 · Cited by 177
Self-citationRedirection of center-of-mass velocity during the step-to-step transition of human walking
Adamczyk, P. G., & Kuo, A. D. (2009) · Journal of Experimental Biology · doi:10.1242/jeb.027581 · Cited by 170
Self-citationEnergetic cost of walking with increased step variability
O’Connor, S. M., Xu, H. Z., & Kuo, A. D. (2012) · Gait & Posture · doi:10.1016/j.gaitpost.2012.01.014 · Cited by 163
Self-citationThe role of series ankle elasticity in bipedal walking
Zelik, K. E., Huang, T.-W. P., Adamczyk, P. G., & Kuo, A. D. (2013) · Journal of Theoretical Biology · doi:10.1016/j.jtbi.2013.12.014 · Cited by 128
Self-citationKowalsky, D. B., Rebula, J. R., Ojeda, L. V., Adamczyk, P. G., & Kuo, A. D. (2021) · PLOS One · doi:10.1371/journal.pone.0228682 · Cited by 106
Self-citationEnergetic cost of producing cyclic muscle force, rather than work, to swing the human leg
Doke, J., & Kuo, A. D. (2007) · Journal of Experimental Biology · doi:10.1242/jeb.02782 · Cited by 104
Self-citationHumans exploit the biomechanics of bipedal gait during visually guided walking over complex terrain
Matthis, J. S., & Fajen, B. R. (2013) · Proceedings of the Royal Society B Biological Sciences · doi:10.1098/rspb.2013.0700 · Cited by 88
Atzler, E., & Herbst, R. (1927) · Pflügers Archiv - European Journal of Physiology · doi:10.1007/bf01731318 · Cited by 71
The high energetic cost of rapid force development in muscle
van der Zee, T. J., & Kuo, A. D. (2021) · Journal of Experimental Biology · doi:10.1242/jeb.233965 · Cited by 47
Self-citationHumans plan for the near future to walk economically on uneven terrain
Darici, O., & Kuo, A. D. (2023) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2211405120 · Cited by 47
Self-citationMahaki, M., Bruijn, S. M., & van Dieën, J. H. (2019) · PeerJ · doi:10.7717/peerj.7939 · Cited by 45
Ojeda, L. V., Rebula, J. R., Kuo, A. D., & Adamczyk, P. G. (2015) · Medical Engineering & Physics · doi:10.1016/j.medengphy.2015.06.010 · Cited by 43
Self-citationDeterminants of preferred ground clearance during swing phase of human walking
Wu, A. R., & Kuo, A. D. (2016) · Journal of Experimental Biology · doi:10.1242/jeb.137356 · Cited by 32
Self-citationUneven terrain treadmill walking in younger and older adults
Downey, R. J., Richer, N., Gupta, R., Liu, C., Pliner, E. M., Roy, A., Hwang, J., Clark, D. J., Hass, C. J., Manini, T. M., et al. (2022) · PLoS ONE · doi:10.1371/journal.pone.0278646 · Cited by 29
Optimal regulation of bipedal walking speed despite an unexpected bump in the road
Darici, O., Temeltaş, H., & Kuo, A. D. (2018) · PLoS ONE · doi:10.1371/journal.pone.0204205 · Cited by 25
Self-citationAnticipatory Control of Momentum for Bipedal Walking on Uneven Terrain
Darici, O., Temeltaş, H., & Kuo, A. D. (2020) · Scientific Reports · doi:10.1038/s41598-019-57156-6 · Cited by 24
Self-citationSoft tissue deformations explain most of the mechanical work variations of human walking
van der Zee, T. J., & Kuo, A. D. (2021) · Journal of Experimental Biology · doi:10.1242/jeb.239889 · Cited by 24
Self-citationBarclay, C. J. (2018) · Elsevier eBooks · doi:10.1016/b978-0-12-814593-7.00006-2 · Cited by 19
Jessup, L. N., Kelly, L. A., Cresswell, A. G., & Lichtwark, G. A. (2023) · Journal of Applied Physiology · doi:10.1152/japplphysiol.00349.2023 · Cited by 11
The metabolic cost of walking balance control and adaptation in young adults
Ahuja, S., & Franz, J. R. (2022) · Gait & Posture · doi:10.1016/j.gaitpost.2022.05.031 · Cited by 10
A model-based analysis of the mechanical cost of walking on uneven terrainPreprint
Voloshina, A. S., Kuo, A. D., Ferris, D. P., & Remy, C. D. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.06.15.152330 · Cited by 9
Self-citationA split-belt instrumented treadmill with uneven terrain
Hosseini-Yazdi, S.-S., & Kuo, A. D. (2024) · Journal of Biomechanics · doi:10.1016/j.jbiomech.2024.112376 · Cited by 4
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Hosseini-Yazdi, S.-S., & Bertram, J. E. A. (2025) · Journal of Theoretical Biology · doi:10.1016/j.jtbi.2025.112234 · Cited by 7
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Metadata
- Authors
- Hosseini-Yazdi, S.-S., & Kuo, A. D.
- Affiliations
- University of Calgary 1 Department of Biomedical Engineering , , Calgary, AB , , T2N 1N4
- ORCID
- Kuo, Arthur D.
- Venue
- Journal of Experimental Biology
- Publisher
- The Company of Biologists
- Published
- 2025-03-15
- Volume
- 228
- Issue
- 6
- ISSN
- 0022-0949 (ISSN-L)
- 1477-9145
- OpenAlex
- W4406963223
- Cited by
- 10
- References
- 49
- Citations by year
- 2026: 6 · 2025: 3
- FWCI
- 1.89
- Type
- journal article
- Language
- en
- Open access
- green
- doi.org
- Licence
- Locations
- Journal of Experimental Biology (publishedVersion)
- bioRxiv (Cold Spring Harbor Laboratory) (submittedVersion, cc-by-nc)
- PubMed (publishedVersion)
- Funders
- Natural Sciences and Engineering Research Council of Canada
- Topics
- Physical Sciences › Engineering › Biomedical Engineering › Lower Extremity Biomechanics and Pathologies
- Health Sciences › Health Professions › Physical Therapy, Sports Therapy and Rehabilitation › Balance, Gait, and Falls Prevention
- Physical Sciences › Engineering › Control and Systems Engineering › Robotic Locomotion and Control
- Keywords
- energy cost of walking, uneven terrain, metabolic rate, positive work, walking speed, pendulum model, walking efficiency, foot clearance, stance phase, gait biomechanics
- MeSH terms
- Adult, Biomechanical Phenomena, Energy Metabolism, Female, Humans, Male, Walking, Young Adult
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