Ankle plantar flexor force production is an important determinant of the preferred walk-to-run transition speed

SUMMARY The mechanisms that govern the voluntary transition from walking to running as walking speed increases in human gait are not well understood. The objective of this study was to examine the hypothesis that plantar flexor muscle force production is greatly impaired at the preferred transition speed (PTS) due to intrinsic muscle properties and, thus, serves as a determinant for the walk-to-run transition. The plantar flexors have been shown to be important contributors to satisfying the mechanical energetic demands of walking and are the primary contributors to the observed ground reaction forces (GRFs) during the propulsion phase. Thus, if the plantar flexor force production begins to diminish near the PTS despite an increase in muscle activation, then a corresponding decrease in the GRFs during the propulsion phase would be expected. This expectation was supported. Both the peak anterior/posterior and vertical GRFs decreased during the propulsion phase at walking speeds near the PTS. A similar decrease was not observed during the braking phase. Further analysis using forward dynamics simulations of walking at increasing speeds and running at the PTS revealed that all lower extremity muscle forces increased with walking speed, except the ankle plantar flexors. Despite an increase in muscle activation with walking speed, the gastrocnemius muscle force decreased with increasing speed, and the soleus force decreased for walking speeds exceeding 80% PTS. These decreases in force production were attributed to the intrinsic force–length–velocity properties of muscle. In addition, the running simulation analysis revealed that the plantar flexor forces nearly doubled for similar activation levels when the gait switched to a run at the PTS due to improved contractile conditions. These results suggest the plantar flexors may serve as an important determinant for the walk-to-run transition and highlight the important role intrinsic muscle properties play in determining the specific neuromotor strategies used in human locomotion.

[1]  HighWire Press Journal of experimental biology , 2022 .

[2]  E. Delagi,et al.  Anatomical guide for the electromyographer : the limbs and trunk /by Edward F. Delagi [et al.] ; illustrated by Phyllis B. Hammond, Aldo O. Perotto, and Hugh Thomas , 2005 .

[3]  A. Thorstensson,et al.  Adaptations to changing speed in human locomotion: speed of transition between walking and running. , 1987, Acta physiologica Scandinavica.

[4]  M L Audu,et al.  A dynamic optimization technique for predicting muscle forces in the swing phase of gait. , 1987, Journal of biomechanics.

[5]  A. Thorstensson,et al.  Ground reaction forces at different speeds of human walking and running. , 1989, Acta physiologica Scandinavica.

[6]  F. Zajac,et al.  A planar model of the knee joint to characterize the knee extensor mechanism. , 1989, Journal of biomechanics.

[7]  F. Zajac Muscle and tendon: properties, models, scaling, and application to biomechanics and motor control. , 1989, Critical reviews in biomedical engineering.

[8]  R. M. Alexander,et al.  Optimization and gaits in the locomotion of vertebrates. , 1989, Physiological reviews.

[9]  F.E. Zajac,et al.  An interactive graphics-based model of the lower extremity to study orthopaedic surgical procedures , 1990, IEEE Transactions on Biomedical Engineering.

[10]  William L. Goffe,et al.  SIMANN: FORTRAN module to perform Global Optimization of Statistical Functions with Simulated Annealing , 1992 .

[11]  Richard A. Brand,et al.  The biomechanics and motor control of human gait: Normal, elderly, and pathological , 1992 .

[12]  S. Simon Gait Analysis, Normal and Pathological Function. , 1993 .

[13]  A Hreljac,et al.  Preferred and energetically optimal gait transition speeds in human locomotion. , 1993, Medicine and science in sports and exercise.

[14]  A Hreljac,et al.  Determinants of the gait transition speed during human locomotion: kinetic factors , 1993 .

[15]  A. Minetti,et al.  The transition between walking and running in humans: metabolic and mechanical aspects at different gradients. , 1994, Acta physiologica Scandinavica.

[16]  S. Olney,et al.  Temporal, kinematic, and kinetic variables related to gait speed in subjects with hemiplegia: a regression approach. , 1994, Physical therapy.

[17]  Michael J. Mueller,et al.  Relationship of plantar-flexor peak torque and dorsiflexion range of motion to kinetic variables during walking. , 1995, Physical therapy.

[18]  S L Delp,et al.  A graphics-based software system to develop and analyze models of musculoskeletal structures. , 1995, Computers in biology and medicine.

[19]  A Hreljac,et al.  Determinants of the gait transition speed during human locomotion: kinematic factors. , 1995, Journal of biomechanics.

[20]  Alan Hreljac,et al.  Effects of physical characteristics on the gait transition speed during human locomotion , 1995 .

[21]  J. Brisswalter,et al.  Energy cost and stride duration variability at preferred transition gait speed between walking and running. , 1996, Canadian journal of applied physiology = Revue canadienne de physiologie appliquee.

[22]  Daniel P. Ferris,et al.  Effect of reduced gravity on the preferred walk-run transition speed. , 1997, The Journal of experimental biology.

[23]  F. Zajac,et al.  Muscle coordination of maximum-speed pedaling. , 1997, Journal of biomechanics.

[24]  A. Minetti,et al.  A theory of metabolic costs for bipedal gaits. , 1997, Journal of theoretical biology.

[25]  W Herzog,et al.  History dependence of force production in skeletal muscle: a proposal for mechanisms. , 1998, Journal of electromyography and kinesiology : official journal of the International Society of Electrophysiological Kinesiology.

[26]  P A Huijing,et al.  Muscle, the motor of movement: properties in function, experiment and modelling. , 1998, Journal of electromyography and kinesiology : official journal of the International Society of Electrophysiological Kinesiology.

[27]  S. Nadeau,et al.  Plantarflexor weakness as a limiting factor of gait speed in stroke subjects and the compensating role of hip flexors. , 1999, Clinical biomechanics.

[28]  R. Kram,et al.  Muscular Force or Work: What Determines the Metabolic Energy Cost of Running? , 2000, Exercise and sport sciences reviews.

[29]  R. R. NEPTUNE,et al.  A Method for Numerical Simulation of Single Limb Ground Contact Events: Application to Heel-Toe Running , 2000, Computer methods in biomechanics and biomedical engineering.

[30]  Robert J. Neal,et al.  Triggers for the transition between human walking and running , 2000 .

[31]  William Anthony Sparrow,et al.  Energetics of human activity , 2000 .

[32]  B I Prilutsky,et al.  Swing- and support-related muscle actions differentially trigger human walk-run and run-walk transitions. , 2001, The Journal of experimental biology.

[33]  F. Zajac,et al.  Contributions of the individual ankle plantar flexors to support, forward progression and swing initiation during walking. , 2001, Journal of biomechanics.

[34]  Richard R Neptune,et al.  Biomechanics and muscle coordination of human walking. Part I: introduction to concepts, power transfer, dynamics and simulations. , 2002, Gait & posture.

[35]  Don W. Morgan,et al.  Comparison between preferred and energetically optimal transition speeds in adolescents , 2002, European Journal of Applied Physiology.

[36]  M. Bobbert,et al.  Mechanics of human triceps surae muscle in walking, running and jumping. , 2002, Acta physiologica Scandinavica.

[37]  A. Minetti,et al.  Biomechanical and physiological aspects of legged locomotion in humans , 2002, European Journal of Applied Physiology.

[38]  Joseph Hamill,et al.  Characteristics of the Vertical Ground Reaction Force Component Prior to Gait Transition , 2002, Research quarterly for exercise and sport.

[39]  B. Abernethy,et al.  Are transitions in human gait determined by mechanical, kinetic or energetic factors? , 2002, Human movement science.

[40]  Richard R Neptune,et al.  Biomechanics and muscle coordination of human walking: part II: lessons from dynamical simulations and clinical implications. , 2003, Gait & posture.

[41]  D. Kerrigan,et al.  Predicting peak kinematic and kinetic parameters from gait speed. , 2003, Gait & posture.

[42]  M. Pandy,et al.  Individual muscle contributions to support in normal walking. , 2003, Gait & posture.

[43]  F. Multon,et al.  Does training have consequences for the walk-run transition speed? , 2003, Human movement science.

[44]  Jacques Mercier,et al.  Energy expenditure and cardiorespiratory responses at the transition between walking and running , 2004, European Journal of Applied Physiology and Occupational Physiology.

[45]  F. Zajac,et al.  Muscle force redistributes segmental power for body progression during walking. , 2004, Gait & posture.

[46]  R R Neptune,et al.  Muscle mechanical work requirements during normal walking: the energetic cost of raising the body's center-of-mass is significant. , 2004, Journal of biomechanics.

[47]  R. Buschbacher Anatomical Guide for the Electromyographer: The Limbs and Trunk , 2007 .