Kinematic patterns while walking on a slope at different speeds.

During walking, the elevation angles of the thigh, shank, and foot (i.e., the angle between the segment and the vertical) covary along a characteristic loop constrained on a plane. Here, we investigate how the shape of the loop and the orientation of the plane, which reflect the intersegmental coordination, change with the slope of the terrain and the speed of progression. Ten subjects walked on an inclined treadmill at different slopes (between -9° and +9°) and speeds (from 0.56 to 2.22 m/s). A principal component analysis was performed on the covariance matrix of the thigh, shank, and foot elevation angles. At each slope and speed, the variance accounted for by the two principal components was >99%, indicating that the planar covariation is maintained. The two principal components can be associated to the limb orientation (PC1*) and the limb length (PC2*). At low walking speeds, changes in the intersegmental coordination across slopes are characterized mainly by a change in the orientation of the covariation plane and in PC2* and to a lesser extent, by a change in PC1*. As speed increases, changes in the intersegmental coordination across slopes are more related to a change in PC1 *, with limited changes in the orientation of the plane and in PC 2*. Our results show that the kinematic patterns highly depend on both slope and speed. NEW & NOTEWORTHY In this paper, changes in the lower-limb intersegmental coordination during walking with slope and speed are linked to changes in the trajectory of the body center of mass. Modifications in the kinematic pattern with slope depend on speed: at slow speeds, the net vertical displacement of the body during each step is related to changes in limb length and orientation. When speed increases, the vertical displacement is mostly related to a change in limb orientation.

[1]  Andy Ruina,et al.  Energetic Consequences of Walking Like an Inverted Pendulum: Step-to-Step Transitions , 2005, Exercise and sport sciences reviews.

[2]  Stacie A. Chvatal,et al.  Review and perspective: neuromechanical considerations for predicting muscle activation patterns for movement , 2012, International journal for numerical methods in biomedical engineering.

[3]  Francesco Lacquaniti,et al.  Patterned control of human locomotion , 2012, The Journal of physiology.

[4]  A. Büschges,et al.  Speed-dependent interplay between local pattern-generating activity and sensory signals during walking in Drosophila , 2016, Journal of Experimental Biology.

[5]  R J Full,et al.  Templates and anchors: neuromechanical hypotheses of legged locomotion on land. , 1999, The Journal of experimental biology.

[6]  Guido Pasquini,et al.  Aging does not affect the intralimb coordination elicited by slip-like perturbation of different intensities. , 2017, Journal of neurophysiology.

[7]  N. L. Svensson,et al.  The influence of surface slope on human gait characteristics: a study of urban pedestrians walking on an inclined surface. , 1996, Ergonomics.

[8]  J. Chow,et al.  Intersegmental coordination scales with gait speed similarly in men and women , 2015, Experimental Brain Research.

[9]  F. Lacquaniti,et al.  Development of pendulum mechanism and kinematic coordination from the first unsupported steps in toddlers , 2004, Journal of Experimental Biology.

[10]  Youkeun K. Oh,et al.  Measurement of lower extremity kinematics and kinetics during valley-shaped slope walking , 2015 .

[11]  Francesco Lacquaniti,et al.  Development of a kinematic coordination pattern in toddler locomotion: planar covariation , 2001, Experimental Brain Research.

[12]  M. Redfern,et al.  Biomechanics of descending ramps , 1997 .

[13]  Ronald M. Harris-Warrick,et al.  Frequency-dependent recruitment of V2a interneurons during fictive locomotion in the mouse spinal cord , 2011, Nature communications.

[14]  Noritaka Kawashima,et al.  Distinct sets of locomotor modules control the speed and modes of human locomotion , 2016, Scientific Reports.

[15]  A. Kuo,et al.  Human walking isn't all hard work: evidence of soft tissue contributions to energy dissipation and return , 2010, Journal of Experimental Biology.

[16]  Jonathan B Dingwell,et al.  Voluntary changes in step width and step length during human walking affect dynamic margins of stability. , 2012, Gait & posture.

[17]  F. Lacquaniti,et al.  Interactions between posture and locomotion: motor patterns in humans walking with bent posture versus erect posture. , 2000, Journal of neurophysiology.

[18]  T. Lu,et al.  Influence of inclination angles on intra- and inter-limb load-sharing during uphill walking. , 2014, Gait & posture.

[19]  K. Kawamura,et al.  Gait analysis of slope walking: a study on step length, stride width, time factors and deviation in the center of pressure. , 1991, Acta medica Okayama.

[20]  Martyna J Grabowska,et al.  Quadrupedal gaits in hexapod animals – inter-leg coordination in free-walking adult stick insects , 2012, Journal of Experimental Biology.

[21]  P. Holmes,et al.  Frontiers in Neural Circuits Neural Circuits , 2022 .

[22]  R. Cham,et al.  Heel contact dynamics during slip events on level and inclined surfaces , 2002 .

[23]  H. Cruse What mechanisms coordinate leg movement in walking arthropods? , 1990, Trends in Neurosciences.

[24]  Noritaka Kawashima,et al.  Speed dependency in α-motoneuron activity and locomotor modules in human locomotion: indirect evidence for phylogenetically conserved spinal circuits , 2017, Proceedings of the Royal Society B: Biological Sciences.

[25]  J. Fetcho,et al.  Spinal Interneurons Differentiate Sequentially from Those Driving the Fastest Swimming Movements in Larval Zebrafish to Those Driving the Slowest Ones , 2009, The Journal of Neuroscience.

[26]  F. Lacquaniti,et al.  Motor Patterns in Walking. , 1999, News in physiological sciences : an international journal of physiology produced jointly by the International Union of Physiological Sciences and the American Physiological Society.

[27]  M. Kuster,et al.  Kinematic and kinetic comparison of downhill and level walking. , 1995, Clinical biomechanics.

[28]  G. W. Lange,et al.  Electromyographic and kinematic analysis of graded treadmill walking and the implications for knee rehabilitation. , 1996, The Journal of orthopaedic and sports physical therapy.

[29]  Shuyang Han,et al.  The Influence of Walking Speed on Gait Patterns During Upslope Walking , 2015 .

[30]  J. Frank,et al.  Locomotor adaptations for changes in the slope of the walking surface. , 2004, Gait & posture.

[31]  Stephen D. Prentice,et al.  Intersegmental coordination while walking up inclined surfaces: age and ramp angle effects , 2008, Experimental Brain Research.

[32]  L C Hunter,et al.  The cost of walking downhill: is the preferred gait energetically optimal? , 2010, Journal of biomechanics.

[33]  B. McFadyen,et al.  Increased Obstacle Clearance in People with ARCA-1 Results in Part from Voluntary Coordination Changes Between the Thigh and Shank Segments , 2011, The Cerebellum.

[34]  Kinematic covariation in pediatric, adult and elderly subjects: is gait control influenced by age? , 2012, Clinical biomechanics.

[35]  Bénédicte Schepens,et al.  Determination of the vertical ground reaction forces acting upon individual limbs during healthy and clinical gait. , 2016, Gait & posture.

[36]  R. M. Alexander Human walking and running , 1984 .

[37]  N. A. Borghese,et al.  Kinematic determinants of human locomotion. , 1996, The Journal of physiology.

[38]  F Lacquaniti,et al.  Locomotor patterns in cerebellar ataxia. , 2014, Journal of neurophysiology.

[39]  P. Willems,et al.  Does an instrumented treadmill correctly measure the ground reaction forces? , 2013, Biology Open.

[40]  R. McN. Alexander,et al.  Tendon elasticity and positional control , 1995, Behavioral and Brain Sciences.

[41]  T. Lu,et al.  Redistribution of intra- and inter-limb support moments during downhill walking on different slopes. , 2014, Journal of biomechanics.

[42]  Marco Schieppati,et al.  Tuning of a basic coordination pattern constructs straight-ahead and curved walking in humans. , 2004, Journal of neurophysiology.

[43]  Irraivan Elamvazuthi,et al.  Biomechanics of Hip, Knee and Ankle joint loading during ascent and descent walking , 2014 .

[44]  C D Mah,et al.  Quantitative analysis of human movement synergies: constructive pattern analysis for gait. , 1994, Journal of motor behavior.

[45]  P. Holmes,et al.  Intra- and intersegmental influences among central pattern generating networks in the walking system of the stick insect. , 2017, Journal of neurophysiology.

[46]  A. McIntosh,et al.  Gait dynamics on an inclined walkway. , 2006, Journal of biomechanics.

[47]  A. d’Avella,et al.  On the origin of planar covariation of elevation angles during human locomotion. , 2008, Journal of neurophysiology.

[48]  J. Schmitz,et al.  Rhythmic patterns in the thoracic nerve cord of the stick insect induced by pilocarpine , 1995, The Journal of experimental biology.

[49]  F. Lacquaniti,et al.  Kinematic coordination in human gait: relation to mechanical energy cost. , 1998, Journal of neurophysiology.

[50]  H. Schwameder,et al.  Effect of sloped walking on lower limb muscle forces. , 2016, Gait & posture.

[51]  P A Willems,et al.  The rebound of the body during uphill and downhill running at different speeds , 2016, Journal of Experimental Biology.

[52]  D. Childress,et al.  Roll-over characteristics of human walking on inclined surfaces. , 2004, Human movement science.

[53]  F. Lacquaniti,et al.  Kinematic control of walking. , 2002, Archives italiennes de biologie.

[54]  P. Willems,et al.  Pendular energy transduction within the step during human walking on slopes at different speeds , 2017, PloS one.

[55]  Andrea N. Lay,et al.  The effects of sloped surfaces on locomotion: a kinematic and kinetic analysis. , 2006, Journal of biomechanics.

[56]  C. T. Farley,et al.  Determinants of the center of mass trajectory in human walking and running. , 1998, The Journal of experimental biology.

[57]  A. Merryweather,et al.  The influence of deformation height on estimating the center of pressure during level and cross-slope walking on sand. , 2015, Gait & posture.

[58]  G. Cavagna,et al.  The sources of external work in level walking and running. , 1976, The Journal of physiology.

[59]  P. Carlson-Kuhta,et al.  Forms of forward quadrupedal locomotion. II. A comparison of posture, hindlimb kinematics, and motor patterns for upslope and level walking. , 1998, Journal of neurophysiology.

[60]  H. Barbeau,et al.  Postural adaptation to walking on inclined surfaces: I. Normal strategies. , 2002, Gait & posture.

[61]  Francesco Lacquaniti,et al.  Modular Control of Limb Movements during Human Locomotion , 2007, The Journal of Neuroscience.

[62]  F. Lacquaniti,et al.  Individual characteristics of human walking mechanics , 1998, Pflügers Archiv.

[63]  Tad McGeer,et al.  Passive Dynamic Walking , 1990, Int. J. Robotics Res..

[64]  T. Higham,et al.  The scaling of uphill and downhill locomotion in legged animals. , 2014, Integrative and comparative biology.

[65]  R. Kram,et al.  Advanced age and the mechanics of uphill walking: a joint-level, inverse dynamic analysis. , 2014, Gait & posture.

[66]  P. Allard,et al.  Effect of trunk inclination on lower limb joint and lumbar moments in able men during the stance phase of gait. , 2009, Clinical biomechanics.

[67]  L Vogt,et al.  Measurement of lumbar spine kinematics in incline treadmill walking. , 1999, Gait & posture.

[68]  A. Büschges,et al.  Inter-leg coordination in the control of walking speed in Drosophila , 2013, Journal of Experimental Biology.

[69]  Francesco Lacquaniti,et al.  Kinematic strategies in newly walking toddlers stepping over different support surfaces. , 2010, Journal of neurophysiology.

[70]  M. Daley,et al.  Compass gait mechanics account for top walking speeds in ducks and humans , 2008, Journal of Experimental Biology.

[71]  Jessica Ausborn,et al.  Separate Microcircuit Modules of Distinct V2a Interneurons and Motoneurons Control the Speed of Locomotion , 2014, Neuron.

[72]  J. Saunders,et al.  The major determinants in normal and pathological gait. , 1953, The Journal of bone and joint surgery. American volume.

[73]  F. Lacquaniti,et al.  Control of foot trajectory in human locomotion: role of ground contact forces in simulated reduced gravity. , 2002, Journal of neurophysiology.

[74]  F. Lacquaniti,et al.  Basal ganglia and gait control: apomorphine administration and internal pallidum stimulation in Parkinson’s disease , 1999, Experimental Brain Research.