Muscle coordination limits efficiency and power output of human limb movement under a wide range of mechanical demands.

This study investigated the influence of cycle frequency and workload on muscle coordination and the ensuing relationship with mechanical efficiency and power output of human limb movement. Eleven trained cyclists completed an array of cycle frequency (cadence)-power output conditions while excitation from 10 leg muscles and power output were recorded. Mechanical efficiency was maximized at increasing cadences for increasing power outputs and corresponded to muscle coordination and muscle fiber type recruitment that minimized both the total muscle excitation across all muscles and the ineffective pedal forces. Also, maximum efficiency was characterized by muscle coordination at the top and bottom of the pedal cycle and progressive excitation through the uniarticulate knee, hip, and ankle muscles. Inefficiencies were characterized by excessive excitation of biarticulate muscles and larger duty cycles. Power output and efficiency were limited by the duration of muscle excitation beyond a critical cadence (120-140 rpm), with larger duty cycles and disproportionate increases in muscle excitation suggesting deteriorating muscle coordination and limitations of the activation-deactivation capabilities. Most muscles displayed systematic phase shifts of the muscle excitation relative to the pedal cycle that were dependent on cadence and, to a lesser extent, power output. Phase shifts were different for each muscle, thereby altering their mechanical contribution to the pedaling action. This study shows that muscle coordination is a key determinant of mechanical efficiency and power output of limb movement across a wide range of mechanical demands and that the excitation and coordination of the muscles is limited at very high cycle frequencies.

[1]  Vinzenz von Tscharner,et al.  Intensity analysis in time-frequency space of surface myoelectric signals by wavelets of specified resolution , 2000 .

[2]  A P Marsh,et al.  Effect of cadence, cycling experience, and aerobic power on delta efficiency during cycling. , 2000, Medicine and science in sports and exercise.

[3]  James M Wakeling,et al.  Muscle coordination during an outdoor cycling time trial. , 2012, Medicine and science in sports and exercise.

[4]  A. Sargeant Structural and functional determinants of human muscle power , 2007, Experimental physiology.

[5]  B. Nigg,et al.  Neuromuscular Strategies during Cycling at Different Muscular Demands. , 2015, Medicine and science in sports and exercise.

[6]  E. Coyle,et al.  Load and Velocity of Contraction Influence Gross and Delta Mechanical Efficiency , 1992, International journal of sports medicine.

[7]  Jostein Hallén,et al.  The most economical cadence increases with increasing workload , 2004, European Journal of Applied Physiology.

[8]  Li Li,et al.  Lower extremity muscle activities during cycling are influenced by load and frequency. , 2003, Journal of electromyography and kinesiology : official journal of the International Society of Electrophysiological Kinesiology.

[9]  D. Sanderson The influence of cadence and power output on the biomechanics of force application during steady-rate cycling in competitive and recreational cyclists. , 1991, Journal of sports sciences.

[10]  David R. Bassett,et al.  The effect of pedaling frequency on glycogen depletion rates in type I and type II quadriceps muscle fibers during submaximal cycling exercise , 2004, European Journal of Applied Physiology and Occupational Physiology.

[11]  J. R. Lacour,et al.  Optimal velocity for maximal power production in non-isokinetic cycling is related to muscle fibre type composition , 2004, European Journal of Applied Physiology and Occupational Physiology.

[12]  M L Hull,et al.  Analysis of EMG measurements during bicycle pedalling. , 1986, Journal of biomechanics.

[13]  M. Ramey,et al.  Influence of pedalling rate and power output on energy expenditure during bicycle ergometry. , 1976, Ergonomics.

[14]  D. Billheimer Functional Data Analysis, 2nd edition edited by J. O. Ramsay and B. W. Silverman , 2007 .

[15]  C. D. De Luca,et al.  Myoelectric signal versus force relationship in different human muscles. , 1983, Journal of applied physiology: respiratory, environmental and exercise physiology.

[16]  B. Silverman,et al.  Functional Data Analysis , 1997 .

[17]  Yvan Champoux,et al.  Interindividual variability of electromyographic patterns and pedal force profiles in trained cyclists , 2008, European Journal of Applied Physiology.

[18]  O. Lippold,et al.  The relation between force and integrated electrical activity in fatigued muscle , 1956, The Journal of physiology.

[19]  D. Farina Counterpoint: spectral properties of the surface EMG do not provide information about motor unit recruitment and muscle fiber type. , 2008, Journal of applied physiology.

[20]  M. Ericson,et al.  On the biomechanics of cycling. A study of joint and muscle load during exercise on the bicycle ergometer. , 1986, Scandinavian journal of rehabilitation medicine. Supplement.

[21]  Vinzenz von Tscharner,et al.  Last word on point:counterpoint: spectral properties of the surface EMG can characterize/do not provide information about motor unit recruitment strategies and muscle fiber type. , 2008, Journal of applied physiology.

[22]  Sylvain Dorel,et al.  Adjustment of muscle coordination during an all-out sprint cycling task. , 2012, Medicine and science in sports and exercise.

[23]  Urs Boutellier,et al.  The generalized force–velocity relationship explains why the preferred pedaling rate of cyclists exceeds the most efficient one , 2005, European Journal of Applied Physiology.

[24]  J. Hagberg,et al.  Effect of pedaling rate on submaximal exercise responses of competitive cyclists. , 1981, Journal of applied physiology: respiratory, environmental and exercise physiology.

[25]  H. Devries,et al.  Mechanomyographic and electromyographic responses during submaximal cycle ergometry , 2000, European Journal of Applied Physiology.

[26]  A. V. van Soest,et al.  Which factors determine the optimal pedaling rate in sprint cycling? , 2000, Medicine and science in sports and exercise.

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

[28]  N. Brown,et al.  Joint-specific power production and fatigue during maximal cycling. , 2009, Journal of biomechanics.

[29]  R R Neptune,et al.  The association between negative muscle work and pedaling rate. , 1999, Journal of biomechanics.

[30]  Dario Farina,et al.  Effect of power, pedal rate, and force on average muscle fiber conduction velocity during cycling. , 2004, Journal of applied physiology.

[31]  Stefano Piazza,et al.  Shared muscle synergies in human walking and cycling. , 2014, Journal of neurophysiology.

[32]  A. Sargeant,et al.  Maximum leg force and power output during short-term dynamic exercise. , 1981, Journal of applied physiology: respiratory, environmental and exercise physiology.

[33]  T. Moritani,et al.  Neuromuscular, metabolic, and kinetic adaptations for skilled pedaling performance in cyclists. , 1998, Medicine and science in sports and exercise.

[34]  A. Guével,et al.  Is interindividual variability of EMG patterns in trained cyclists related to different muscle synergies? , 2010, Journal of applied physiology.

[35]  James M Wakeling,et al.  Muscle coordination patterns for efficient cycling. , 2012, Medicine and science in sports and exercise.

[36]  R. Lepers,et al.  Neuromuscular function during prolonged pedalling exercise at different cadences. , 2005, Acta physiologica Scandinavica.

[37]  R. Patterson,et al.  The influence of flywheel weight and pedalling frequency on the biomechanics and physiological responses to bicycle exercise. , 1983, Ergonomics.

[38]  A J Sargeant,et al.  Human Power Output and Muscle Fatigue , 1994, International journal of sports medicine.

[39]  R. Neptune,et al.  The effect of pedaling rate on coordination in cycling. , 1997, Journal of biomechanics.

[40]  James M Wakeling,et al.  Patterns of motor recruitment can be determined using surface EMG. , 2009, Journal of electromyography and kinesiology : official journal of the International Society of Electrophysiological Kinesiology.

[41]  T. Moritani,et al.  Optimal pedaling rate estimated from neuromuscular fatigue for cyclists. , 1996, Medicine and science in sports and exercise.

[42]  P. D. di Prampero Cycling on Earth, in space, on the Moon , 2000, European journal of applied physiology.

[43]  R. Josephson Dissecting muscle power output. , 1999, The Journal of experimental biology.

[44]  J. A. L. Calbet,et al.  Cycling efficiency and pedalling frequency in road cyclists , 1999, European Journal of Applied Physiology and Occupational Physiology.

[45]  Gertjan Ettema,et al.  Efficiency in cycling: a review , 2009, European Journal of Applied Physiology.

[46]  G. Millet,et al.  Influence of cycling cadence on neuromuscular activity of the knee extensors in humans , 2002, European Journal of Applied Physiology.

[47]  Philip E. Martin,et al.  The relationship between cadence and lower extremity EMG in cyclists and noncyclists. , 1995, Medicine and science in sports and exercise.

[48]  J. Coast,et al.  Linear increase in optimal pedal rate with increased power output in cycle ergometry , 1985, European Journal of Applied Physiology and Occupational Physiology.

[49]  R. Lepers,et al.  Cycling exercise and the determination of electromechanical delay. , 2007, Journal of electromyography and kinesiology : official journal of the International Society of Electrophysiological Kinesiology.

[50]  Gertjan Ettema,et al.  Pedaling technique and energy cost in cycling. , 2011, Medicine and science in sports and exercise.

[51]  J. Wakeling,et al.  Estimating changes in metabolic power from EMG , 2013, SpringerPlus.

[52]  James M. Wakeling,et al.  Early deactivation of slower muscle fibres at high movement frequencies , 2014, Journal of Experimental Biology.

[53]  F. Hug,et al.  Force-velocity relationship in cycling revisited: benefit of two-dimensional pedal forces analysis. , 2009, Medicine and science in sports and exercise.

[54]  David Bendahan,et al.  Heterogeneity of muscle recruitment pattern during pedaling in professional road cyclists: a magnetic resonance imaging and electromyography study , 2004, European Journal of Applied Physiology.

[55]  James M Wakeling,et al.  Neuromechanics of muscle synergies during cycling. , 2009, Journal of neurophysiology.

[56]  Alejandro Lucia,et al.  In professional road cyclists, low pedaling cadences are less efficient. , 2004, Medicine and science in sports and exercise.

[57]  Pierre Samozino,et al.  Why does power output decrease at high pedaling rates during sprint cycling? , 2007, Medicine and science in sports and exercise.

[58]  James M Wakeling,et al.  Spectral properties of myoelectric signals from different motor units in the leg extensor muscles , 2004, Journal of Experimental Biology.

[59]  A. Beelen,et al.  Effect of fatigue on maximal power output at different contraction velocities in humans. , 1991, Journal of applied physiology.

[60]  B. Nigg,et al.  Point: spectral properties of the surface EMG can characterize/do not provide information about motor unit recruitment strategies and muscle fiber type. , 2008, Journal of applied physiology.

[61]  M O Ericson,et al.  Muscular activity during ergometer cycling. , 1985, Scandinavian journal of rehabilitation medicine.

[62]  G Sjøgaard,et al.  Relationship between efficiency and pedal rate in cycling: significance of internal power and muscle fiber type composition , 2006, Scandinavian journal of medicine & science in sports.

[63]  Silvia Conforto,et al.  Inter-individual variability of forces and modular muscle coordination in cycling: a study on untrained subjects. , 2013, Human movement science.

[64]  R. R. Neptune,et al.  Muscle Activation and Deactivation Dynamics: The Governing Properties in Fast Cyclical Human Movement Performance? , 2001, Exercise and sport sciences reviews.

[65]  R R Neptune,et al.  Cadence, power, and muscle activation in cycle ergometry. , 2000, Medicine and science in sports and exercise.

[66]  J. Moreno,et al.  trained cyclists related to different muscle synergies? Is interindividual variability of EMG patterns in , 2015 .

[67]  G. Terzis,et al.  Muscle fibre type composition and body composition in hammer throwers. , 2010, Journal of sports science & medicine.

[68]  R. Marsh,et al.  Optimal shortening velocity (V/Vmax) of skeletal muscle during cyclical contractions: length-force effects and velocity-dependent activation and deactivation. , 1998, The Journal of experimental biology.

[69]  J. Wakeling,et al.  Muscle coordination is key to the power output and mechanical efficiency of limb movements , 2010, Journal of Experimental Biology.

[70]  G. J. van Ingen Schenau,et al.  The constrained control of force and position in multi-joint movements , 1992, Neuroscience.

[71]  J. S. Petrofsky,et al.  Frequency and amplitude analysis of the EMG during exercise on the bicycle ergometer , 1979, European Journal of Applied Physiology and Occupational Physiology.

[72]  Hannover,et al.  Relationship Between Work Load, Pedal Frequency, and Physical Fitness* , 1984, International journal of sports medicine.

[73]  Sabrina S. M. Lee,et al.  Movement mechanics as a determinate of muscle structure, recruitment and coordination , 2011, Philosophical Transactions of the Royal Society B: Biological Sciences.

[74]  M L Hull,et al.  A theoretical basis for interpreting the force applied to the pedal in cycling. , 1993, Journal of biomechanics.

[75]  J. Wakeling,et al.  Muscle fibre recruitment can respond to the mechanics of the muscle contraction , 2006, Journal of The Royal Society Interface.