Spinal cord circuits: are they mirrors of musculoskeletal mechanics?

Over the past decade, research at three different levels of sensorimotor control has revealed a degree of complexity that challenges traditional hypotheses regarding servocontrol of individual muscles: (a) The connectivity of spinal circuits is much more divergent and convergent than expected. (b) The normal and reflex-induced recruitment of individual muscles and compartments of muscles is more finely controlled than was noted previously. (c) The mechanical interactions among linked skeletal segments and their often multiarticular muscles are neither simple nor intuitively obvious. We have developed a mathematical model of the cat hind limb that permits us to examine the influence of individual muscles on posture and gait. We have used linear quadratic control theory to predict the optimal distribution of feedback from a hypothetical set of proprioceptors, given different assumptions about the behavioral goals of the animal. The changes in these predictions that result from changes in the structure and control objectives of the model may provide insights into the functions actually performed by the various circuits in the spinal cord.

[1]  G. C. Huber,et al.  On the form and arrangement in fasciculi of striated voluntary muscle fibers. A preliminary report , 1916 .

[2]  J. Manter The Dynamics Of Quadrupedal Walking , 1938 .

[3]  A. Hill The heat of shortening and the dynamic constants of muscle , 1938 .

[4]  H. Elftman Biomechanics of muscle with particular application to studies of gait. , 1966, The Journal of bone and joint surgery. American volume.

[5]  A. Huxley,et al.  The variation in isometric tension with sarcomere length in vertebrate muscle fibres , 1966, The Journal of physiology.

[6]  Michael Athans,et al.  Optimal Control , 1966 .

[7]  M Abrahams,et al.  Mechanical behaviour of tendon in vitro. A preliminary report. , 1967, Medical & biological engineering.

[8]  A. Bahler,et al.  Modeling of mammalian skeletal muscle. , 1968, IEEE transactions on bio-medical engineering.

[9]  Arthur E. Bryson,et al.  Applied Optimal Control , 1969 .

[10]  G. C. Joyce,et al.  The mechanical properties of cat soleus muscle during controlled lengthening and shortening movements , 1969, The Journal of physiology.

[11]  G. C. Joyce,et al.  Isotonic lengthening and shortening movements of cat soleus muscle , 1969, The Journal of physiology.

[12]  C. P. Taylor,et al.  Isometric muscle contraction and the active state: an analog computer study. , 1969, Biophysical journal.

[13]  K. Edman,et al.  The rising phase of the active state in single skeletal muscle fibres of the frog. , 1970, Acta physiologica Scandinavica.

[14]  M. A. Athans,et al.  The role and use of the stochastic linear-quadratic-Gaussian problem in control system design , 1971 .

[15]  A. Huxley,et al.  Proposed Mechanism of Force Generation in Striated Muscle , 1971, Nature.

[16]  M. Tokuriki 四走獣の歩行運動の筋電図学的・関節機構学的研究 : I. 常歩 , 1973 .

[17]  F. Julian,et al.  Regulation of Force and Speed of Shortening in Muscle Contraction , 1973 .

[18]  G. E. Goslow,et al.  The cat step cycle: Hind limb joint angles and muscle lengths during unrestrained locomotion , 1973, Journal of morphology.

[19]  R. J. Podolsky,et al.  Muscle Contraction Transients, Cross-Bridge Kinetics, and the Fenn Effect , 1973 .

[20]  J. Thorson,et al.  The kinetics of muscle contraction , 1975 .

[21]  H. Hatze,et al.  A complete set of control equations for the human musculo-skeletal system. , 1977, Journal of biomechanics.

[22]  T. Moritani,et al.  Reexamination of the relationship between the surface integrated electromyogram (IEMG) and force of isometric contraction. , 1978, American journal of physical medicine.

[23]  J. G. Andrews,et al.  A biomechanical investigation of the human hip. , 1978, Journal of biomechanics.

[24]  J C Houk,et al.  Regulation of stiffness by skeletomotor reflexes. , 1979, Annual review of physiology.

[25]  At Hof,et al.  EMG to force processing , 1980 .

[26]  J. van den Berg,et al.  EMG to force processing I: An electrical analogue of the Hill muscle model. , 1981, Journal of biomechanics.

[27]  S R Simon,et al.  An evaluation of the approaches of optimization models in the prediction of muscle forces during human gait. , 1981, Journal of biomechanics.

[28]  J. E. Wood,et al.  A sliding-filament cross-bridge ensemble model of muscle contraction for mechanical transients , 1981 .

[29]  J. van den Berg,et al.  EMG to force processing IV: Eccentric-concentric contractions on a spring-flywheel set up. , 1981, Journal of biomechanics.

[30]  G. Goldspink Design of muscle for locomotion and the maintenance of posture , 1981, Trends in Neurosciences.

[31]  F. Zajac,et al.  Hindlimb muscular activity, kinetics and kinematics of cats jumping to their maximum achievable heights. , 1981, The Journal of experimental biology.

[32]  R. F. Ker Dynamic tensile properties of the plantaris tendon of sheep (Ovis aries). , 1981, The Journal of experimental biology.

[33]  S R Simon,et al.  Analysis and synthesis of human swing leg motion during gait and its clinical applications. , 1981, Journal of biomechanics.

[34]  R. Crowninshield,et al.  A physiologically based criterion of muscle force prediction in locomotion. , 1981, Journal of biomechanics.

[35]  J. van den Berg,et al.  EMG to force processing II: Estimation of parameters of the Hill muscle model for the human triceps surae by means of a calfergometer. , 1981, Journal of biomechanics.

[36]  J. van den Berg,et al.  EMG to force processing III: Estimation of model parameters for the human triceps surae muscle and assessment of the accuracy by means of a torque plate. , 1981, Journal of biomechanics.

[37]  F M van Krieken,et al.  A model of lower extremity muscular anatomy. , 1982, Journal of biomechanical engineering.

[38]  C. Gans Fiber architecture and muscle function. , 1982, Exercise and sport sciences reviews.

[39]  R. Roy,et al.  Architecture of the hind limb muscles of cats: Functional significance , 1982, Journal of morphology.

[40]  R W Baildon,et al.  A new approach to the human muscle model. , 1983, Journal of biomechanics.

[41]  G H Pollack,et al.  The cross-bridge theory. , 1983, Physiological reviews.

[42]  William S. Levine,et al.  Neuromuscular and Musculoskeletal Control Models for the Human Leg , 1983 .

[43]  J. Halbertsma The stride cycle of the cat: the modelling of locomotion by computerized analysis of automatic recordings. , 1983, Acta physiologica Scandinavica. Supplementum.

[44]  H Hemami,et al.  Control of sliding and rolling at natural joints. , 1984, Journal of biomechanical engineering.

[45]  N. Hogan Adaptive control of mechanical impedance by coactivation of antagonist muscles , 1984 .

[46]  Yuan F. Zheng,et al.  Muscle Dynamics, Size Principle, and Stability , 1984, IEEE Transactions on Biomedical Engineering.

[47]  D. Winter,et al.  Predictions of knee and ankle moments of force in walking from EMG and kinematic data. , 1985, Journal of biomechanics.

[48]  Gerald E. Loeb,et al.  Optimal control principles for sensory transducers , 1985 .

[49]  Ronald F. Zernicke,et al.  Modulation of limb dynamics in the swing phase of locomotion , 1985 .

[50]  Michael R. Pierrynowski,et al.  Estimating the muscle forces generated in the human lower extremity when walking: a physiological solution , 1985 .

[51]  Michael R. Pierrynowski,et al.  A physiological model for the evaluation of muscular forces in human locomotion: theoretical aspects , 1985 .

[52]  R. Cooke,et al.  The mechanism of muscle contraction. , 1986, CRC critical reviews in biochemistry.

[53]  D. McCrea Spinal cord circuitry and motor reflexes. , 1986, Exercise and sport sciences reviews.

[54]  O. I. Fukson,et al.  Adaptability of innate motor patterns and motor control mechanisms , 1986, Behavioral and Brain Sciences.

[55]  G. Loeb,et al.  Electromyography for Experimentalists , 1986 .

[56]  D. McCloskey,et al.  The role of joint receptors in human kinaesthesia when intramuscular receptors cannot contribute. , 1987, The Journal of physiology.

[57]  G. Stein,et al.  The LQG/LTR procedure for multivariable feedback control design , 1987 .

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

[59]  J. Macpherson Strategies that simplify the control of quadrupedal stance. I. Forces at the ground. , 1988, Journal of neurophysiology.

[60]  J M Macpherson,et al.  Strategies that simplify the control of quadrupedal stance. II. Electromyographic activity. , 1988, Journal of neurophysiology.

[61]  E. Otten Concepts and Models of Functional Architecture in Skeletal Muscle , 1988, Exercise and sport sciences reviews.

[62]  William S. Levine,et al.  The modeling of the neuro-musculo-skeletal control system of a cat hindlimb , 1988, Proceedings IEEE International Symposium on Intelligent Control 1988.

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