The motor system controls what it senses

marshalling the free variables into coherent functions. Quadruped gait may be an example: when one stable movement pattern is driven beyond a critical value on a system-sensitive parameter, a bifurcation occurs and a new spatiotemporal pattern a new stability arises. In such a view, no explicit "gait selection" process is required (e.g., Gallistel 1980). To reiterate our main point, however, in the perspective offered here, order (control and coordination) is functionally specified in the system's dynamics. The radical claim, as Gibson (1979) once remarked, is that behavior is regular but there are no regulators. A less radical statement would be an affirmative answer to Yates's (1980) question to the readers of the American Journal of Physiology: Do [you] know of a serious effort to discharge the homunculus? The spirit of the foregoing discussion leads us to raise one final issue. It is the growing intuition stemming from theoretical considerations raised here and elsewhere (cf. Anderson 1972) that the problem of order in natural systems might be attacked more effectively by seeking out a single set of physical principles that can apply at all levels, rather than by positing different units of analysis at each level. One assumes that nature operates with ancient themes. In this commentary we have tried to provide the flavor of the ones that neuroscience in general, and the field of motor control in particular, might consider worth orchestrating.

[1]  J. Eccles,et al.  The isometric responses of mammalian muscles , 1930, The Journal of physiology.

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

[3]  E. Asmussen,et al.  Body Temperature and Capacity for Work , 1945 .

[4]  A. Hill The abrupt transition from rest to activity in muscle , 1949, Proceedings of the Royal Society of London. Series B - Biological Sciences.

[5]  R. Granit,et al.  Influence of stimulation of central nervous structures on muscle spindles in cat. , 1953, Acta physiologica Scandinavica.

[6]  L. Cohen Localization of stretch reflex. , 1953, Journal of neurophysiology.

[7]  L. Cohen Organization of stretch reflex into two types of direct spinal arcs. , 1954, Journal of neurophysiology.

[8]  G. A. Miller THE PSYCHOLOGICAL REVIEW THE MAGICAL NUMBER SEVEN, PLUS OR MINUS TWO: SOME LIMITS ON OUR CAPACITY FOR PROCESSING INFORMATION 1 , 1956 .

[9]  P. Matthews A study of certain factors influencing the stretch reflex of the decerebrate cat , 1959, The Journal of physiology.

[10]  P. Matthews The dependence of tension upon extension in the stretch reflex of the soleus muscle of the decerebrate cat , 1959, The Journal of physiology.

[11]  O J SMITH,et al.  Nonlinear computations in the human controller. , 1962, IRE transactions on bio-medical electronics.

[12]  George A. Bekey,et al.  The Human Operator as a Sampled-Data System , 1962 .

[13]  T. Roberts RHYTHMIC EXCITATION OF A STRETCH REFLEX, REVEALING (A) HYSTERESIS AND (B) A DIFFERENCE BETWEEN THE RESPONSES TO PULLING AND TO STRETCHING. , 1963, Quarterly journal of experimental physiology and cognate medical sciences.

[14]  L. Stark,et al.  A Discrete Model for Eye Tracking Movements , 1963, IEEE Transactions on Military Electronics.

[15]  Laurence R. Young,et al.  Variable Feedback Experiments Testing a Sampled Data Model for Eye Tracking Movements , 1963 .

[16]  P. Matthews MUSCLE SPINDLES AND THEIR MOTOR CONTROL. , 1964, Physiological reviews.

[17]  E. Henneman,et al.  RELATIONS BETWEEN STRUCTURE AND FUNCTION IN THE DESIGN OF SKELETAL MUSCLES. , 1965, Journal of neurophysiology.

[18]  G. Eklund,et al.  EXTERNAL INTERCOSTAL AND PHRENIC ALPHA-MOTOR RESPONSES TO CHANGES IN RESPIRATORY LOAD. , 1965, Acta physiologica Scandinavica.

[19]  D. Kernell The Limits of Firing Frequency in Cat Lumbosacral Motoneurones Possessing Different Time Course of Afterhyperpolarization , 1965 .

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

[21]  J. Houk,et al.  Responses of Golgi tendon organs to active contractions of the soleus muscle of the cat. , 1967, Journal of neurophysiology.

[22]  M. Bárány,et al.  ATPase Activity of Myosin Correlated with Speed of Muscle Shortening , 1967, The Journal of general physiology.

[23]  E. Evarts,et al.  Relation of pyramidal tract activity to force exerted during voluntary movement. , 1968, Journal of neurophysiology.

[24]  L. Stark,et al.  Sampling or intermittency in hand control system dynamics. , 1968, Biophysical journal.

[25]  L Stark,et al.  The human eye-movement mechanism. Experiments, modeling, and model testing. , 1968, Archives of ophthalmology.

[26]  Duane T. McRuer,et al.  A Neuromuscular Actuation System Model , 1968 .

[27]  P. Matthews,et al.  The sensitivity of muscle spindle afferents to small sinusoidal changes of length , 1969, The Journal of physiology.

[28]  P. Matthews Evidence that the secondary as well as the primary endings of the muscle spindles may be responsible for the tonic stretch reflex of the decerebrate cat , 1969, The Journal of physiology.

[29]  H. Táboříková,et al.  Conditioning of H-reflexes by a preceding subthreshold H-reflex stimulus. , 1969, Brain : a journal of neurology.

[30]  Laurence R. Young,et al.  The current status of vestibular system models , 1969, Autom..

[31]  J B Morrison,et al.  The mechanics of muscle function in locomotion. , 1970, Journal of biomechanics.

[32]  L. Stark,et al.  Interactions between voluntary and postural mechanisms of thehuman motor system. , 1970, Journal of neurophysiology.

[33]  J. Houk,et al.  An evaluation of length and force feedback to soleus muscles of decerebrate cats. , 1970, Journal of neurophysiology.

[34]  P. MacNeilage Motor control of serial ordering of speech. , 1970, Psychological review.

[35]  D R Humphrey,et al.  Predicting Measures of Motor Performance from Multiple Cortical Spike Trains , 1970, Science.

[36]  S. Grillner,et al.  Motor unit activity and stiffness of the contracting muscle fibres in the tonic stretch reflex. , 1971, Acta physiologica Scandinavica.

[37]  R. Almond The therapeutic community. , 1971, Scientific American.

[38]  L. Stark THE CONTROL SYSTEM FOR VERSIONAL EYE MOVEMENTS , 1971 .

[39]  D. Humphrey,et al.  Relating motor cortex spike trains to measures of motor performance. , 1972, Brain research.

[40]  Easton Ta On the normal use of reflexes. , 1972 .

[41]  K. Pearson Central Programming and Reflex Control of Walking in the Cockroach , 1972 .

[42]  P. Anderson More is different. , 1972, Science.

[43]  G. Inbar Muscle spindles in muscle control , 1972, Kybernetik.

[44]  C. D. MARSDEN,et al.  Servo Action in Human Voluntary Movement , 1972, Nature.

[45]  L. D. Partridge,et al.  Interrelationships studied in a semibiological "reflex". , 1972, The American journal of physiology.

[46]  A. G. Feldman,et al.  The influence of different descending systems on the tonic stretch reflex in the cat. , 1972, Experimental neurology.

[47]  J. Abbs,et al.  A Two‐Dimensional Strain Gauge Transducer System for the Lips and Jaw , 1973 .

[48]  R. Stein,et al.  Determination of the frequency response of isometric soleus muscle in the cat using random nerve stimulation , 1973, The Journal of physiology.

[49]  G. E. Goslow,et al.  Time Constraints for Inter-Limb Co-Ordination in the Cat During Unrestrained Locomotion , 1973 .

[50]  T. Nichols Reflex and Non-Reflex Stiffness of Soleus Muscle in the Cat , 1973 .

[51]  A. Vallbo,et al.  The Significance of Intramuscular Receptors in Load Compensation During Voluntary Contractions in Man , 1973 .

[52]  T. Sears,et al.  Monosynaptic excitation of motoneurones from secondary endings of muscle spindles , 1974, Nature.

[53]  R. Stein Peripheral control of movement. , 1974, Physiological reviews.

[54]  M. Goldberger,et al.  Restitution of function and collateral sprouting in the cat spinal cord: The deafferented animal , 1974, The Journal of comparative neurology.

[55]  V. Edgerton,et al.  Motor Unit Properties and Selective Involvement In Movement , 1975, Exercise and sport sciences reviews.

[56]  J. Parlange,et al.  Water uptake, diameter change, and nonlinear diffusion in tree stems. , 1975, Plant physiology.

[57]  L. Stark,et al.  Time optimal behavior of human saccadic eye movement , 1975 .

[58]  V. V. Krishnan,et al.  Inners and biocontrol models. , 1975, Bulletin of mathematical biology.

[59]  S. Grillner Locomotion in vertebrates: central mechanisms and reflex interaction. , 1975, Physiological reviews.

[60]  Bruce Bridgeman,et al.  Failure to detect displacement of the visual world during saccadic eye movements , 1975, Vision Research.

[61]  F. Richmond,et al.  Morphology and distribution of muscle spindles in dorsal muscles of the cat neck. , 1975, Journal of neurophysiology.

[62]  F. Zajac,et al.  The effect of activation history on tension production by individual muscle units , 1976, Brain Research.

[63]  G. Jones,et al.  An adaptive neural model compatible with plastic changes induced in the human vestibulo-ocular reflex by prolonged optical reversal of vision , 1976, Brain Research.

[64]  M. L. Shik,et al.  Neurophysiology of locomotor automatism. , 1976, Physiological reviews.

[65]  E. Bizzi,et al.  Mechanisms underlying achievement of final head position. , 1976, Journal of neurophysiology.

[66]  G F Inbar,et al.  Parameter and signal adaptation in the stretch reflex loop. , 1976, Progress in brain research.

[67]  H. Hatze The complete optimization of a human motion , 1976 .

[68]  G. M. Goodwin The Sense Of Limb Position And Movement , 1976, Exercise and sport sciences reviews.

[69]  J. Houk,et al.  Improvement in linearity and regulation of stiffness that results from actions of stretch reflex. , 1976, Journal of neurophysiology.

[70]  P. Milgram,et al.  Distortion Suppression in Neuromuscular Information Transmission Due to Interchannel Dispersion in Muscle Spindle Firing Thresholds , 1976, IEEE Transactions on Biomedical Engineering.

[71]  Stark Lawrence,et al.  Saccadic suppression of image displacement , 1976, Vision Research.

[72]  R. Reinking,et al.  Analysis of muscle receptor connections by spike-triggered averaging. 2. Spindle group II afferents. , 1976, Journal of neurophysiology.

[73]  J. Abbs,et al.  Labial-Mandibular Coordination in the Production of Speech: Implications for the Operation of Motor Equivalence , 1976, Phonetica.

[74]  J H Abbs,et al.  Additional observations on responses to resistive loading of the jaw. , 1976, Journal of speech and hearing research.

[75]  D. Robinson Adaptive gain control of vestibuloocular reflex by the cerebellum. , 1976, Journal of neurophysiology.

[76]  J. Houk,et al.  Regulatory actions of human stretch reflex. , 1976, Journal of neurophysiology.

[77]  V. Edgerton,et al.  EMG of slow and fast ankle extensors of cat during posture, locomotion, and jumping. , 1977, Journal of neurophysiology.

[78]  A. Iberall A field and circuit thermodynamics for integrative physiology. I. Introduction to the general notions. , 1977, The American journal of physiology.

[79]  J. Kelso Motor control mechanisms underlying human movement reproduction. , 1977, Journal of experimental psychology. Human perception and performance.

[80]  K. Stevens Physics of Laryngeal Behavior and Larynx Modes , 1977, Phonetica.

[81]  A Iberall,et al.  A global model of neuronal command-control systems. , 1977, Bio Systems.

[82]  W. Gonyea,et al.  Morphological and histochemical organization of the flexor carpi radialis muscle in the cat. , 1977, The American journal of anatomy.

[83]  A. Iberall,et al.  Homeokinetics: A Physical Science for Complex Systems , 1978, Science.

[84]  D. Stuart,et al.  Functional anatomy of the association between motor units and muscle receptors , 1978 .

[85]  P. Roland Sensory feedback to the cerebral cortex during voluntary movement in man. , 1978 .

[86]  J. A. Scott Kelso,et al.  Changing views of feedforward and feedback in voluntary movement , 1978, Behavioral and Brain Sciences.

[87]  Motohide Umano,et al.  FSTDS System: A fuzzy-set manipulation system , 1978, Inf. Sci..

[88]  E. Bizzi,et al.  Processes controlling arm movements in monkeys. , 1978, Science.

[89]  David E. Hardt,et al.  Determining Muscle Forces in the Leg During Normal Human Walking—An Application and Evaluation of Optimization Methods , 1978 .

[90]  A. Iberall A field and circuit thermodynamics for integrative physiology. III. Keeping the books--a general experimental method. , 1978, The American journal of physiology.

[91]  János Szentágothai,et al.  A false alternative , 1978, Behavioral and Brain Sciences.

[92]  B. Walmsley,et al.  Forces produced by medial gastrocnemius and soleus muscles during locomotion in freely moving cats. , 1978, Journal of neurophysiology.

[93]  M. D. Neilson,et al.  The role of action reflexes in the damping of mechanical oscillations , 1978, Brain Research.

[94]  Roy D. Crowninshield,et al.  Use of Optimization Techniques to Predict Muscle Forces , 1978 .

[95]  Antonio Pedotti,et al.  Optimization of muscle-force sequencing in human locomotion , 1978 .

[96]  E. Bizzi,et al.  Effect of load disturbances during centrally initiated movements. , 1978, Journal of neurophysiology.

[97]  E. Luschei,et al.  The strength of the reflex response to sinusoidal stretch of monkey jaw closing muscles during voluntary contraction. , 1978, The Journal of physiology.

[98]  F. A. Miles,et al.  Concepts of motor organization. , 1979, Annual review of psychology.

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

[100]  J. M. Hofferer,et al.  Visual-motor adaptation. Quantitative demonstration in patients with posterior fossa involvement. , 1979, Archives of neurology.

[101]  J. F. Soechting,et al.  Reflex motor output to torque pulses in man: Identification of short- and long-latency loops with individual feedback parameters , 1979, Neuroscience.

[102]  R. Stein,et al.  Optimal motor patterns for activating mammalian muscle , 1979, Brain Research.

[103]  E. Bizzi,et al.  Characteristics of motor programs underlying arm movements in monkeys. , 1979, Journal of neurophysiology.

[104]  J. Duysens,et al.  Activity patterns in individual hindlimb primary and secondary muscle spindle afferents during normal movements in unrestrained cats. , 1979, Journal of neurophysiology.

[105]  P. Neilson,et al.  Tonic stretch reflexes in lip, tongue and jaw muscles , 1979, Brain Research.

[106]  A. Prochazka,et al.  Muscle spindle discharge in normal and obstructed movements. , 1979, The Journal of physiology.

[107]  Elliot Saltzman,et al.  Levels of sensorimotor representation , 1979 .

[108]  D Goodman,et al.  On the nature of human interlimb coordination. , 1979, Science.

[109]  P. Rudomín,et al.  The influence of the gamma system on cross-correlated activity of Ia muscle spindles and its relation to information transmission , 1979, Neuroscience Letters.

[110]  W. Becker,et al.  An analysis of the saccadic system by means of double step stimuli , 1979, Vision Research.

[111]  J. Cooke The role of stretch reflexes during active movements , 1980, Brain Research.

[112]  V. Edgerton,et al.  Rapid ankle extension during paw shakes: selective recruitment of fast ankle extensors. , 1980, Journal of neurophysiology.

[113]  Richard A. Schmidt,et al.  Terminal Accuracy of Unexpectedly Loaded Rapid Movements , 1980 .

[114]  F. Delcomyn Neural basis of rhythmic behavior in animals. , 1980, Science.

[115]  E. Luschei,et al.  Evidence that the human jaw stretch reflex increases the resistance of the mandible to small displacements. , 1980, The Journal of physiology.

[116]  R. Schmidt Past and future issues in motor programming. , 1980, Research quarterly for exercise and sport.

[117]  J. Cooke 11 The Organization of Simple, Skilled Movements , 1980 .

[118]  K. Appenteng,et al.  Fusimotor activity in masseter nerve of the cat during reflex jaw movements. , 1980, The Journal of physiology.

[119]  J. F. Soechting,et al.  Modulation of the myotatic reflex gain in man during intentional movements , 1980, Brain Research.

[120]  R. Stein Nerve and Muscle: Membranes, Cells, and Systems , 1980 .

[121]  A. G. Feldman Superposition of motor programs—II. Rapid forearm flexion in man , 1980, Neuroscience.

[122]  G. Gottlieb,et al.  Response to sudden torques about ankle in man. III. Suppression of stretch-evoked responses during phasic contraction. , 1980, Journal of neurophysiology.

[123]  J. Lanman Movement and the mechanical properties of the intact human elbow joint. , 1980 .

[124]  A. G. Feldman Superposition of motor programs—I. Rhythmic forearm movements in man , 1980, Neuroscience.

[125]  P. Howell,et al.  Isotonic vocalis contraction as a means of producing rapid decreases in Fo. , 1980, Journal of speech and hearing research.

[126]  W. Rymer,et al.  Absence of force-feedback regulation in soleus muscle of the decerebrate cat , 1980, Brain Research.

[127]  J. Kelso,et al.  Exploring a vibratory systems analysis of human movement production. , 1980, Journal of neurophysiology.

[128]  E. Bizzi 7 Central and Peripheral Mechanisms in Motor Control , 1980 .

[129]  Duane T. McRuer,et al.  Human dynamics in man-machine systems , 1980, Autom..

[130]  E. Fetz,et al.  Functional classes of primate corticomotoneuronal cells and their relation to active force. , 1980, Journal of neurophysiology.

[131]  H. Cruse,et al.  Is the position of the femur tibia joint under feedback control in the walking stick insect?: II. Electrophysiological recordings , 1981 .

[132]  F Lacquaniti,et al.  Time-varying properties of myotatic response in man during some simple motor tasks. , 1981, Journal of neurophysiology.

[133]  R. Reinking,et al.  "Sensory partitioning" of cat medial gastrocnemius muscle by its muscle spindles and tendon organs. , 1981, Journal of neurophysiology.

[134]  J. Houk,et al.  Dependence of dynamic response of spindle receptors on muscle length and velocity. , 1981, Journal of neurophysiology.

[135]  H. Johansson,et al.  Reflex activation of dynamic fusimotor neurons by natural stimulation of muscle and joint receptor afferent units , 1981 .

[136]  Michael J. O'Donovan,et al.  Discharge patterns of hindlimb motoneurons during normal cat locomotion. , 1981, Science.

[137]  R. Stein,et al.  Nonlinear summation of contractions in cat muscles. II. Later facilitation and stiffness changes , 1981, The Journal of general physiology.

[138]  V. C. Abrahams Comment: unresolved questions concerning muscle afferents. , 1981, Canadian journal of physiology and pharmacology.

[139]  C. Marsden,et al.  Reliability and efficacy of the long‐latency stretch reflex in the human thumb. , 1981, The Journal of physiology.

[140]  Holk Cruse,et al.  Is the position of the femur tibia joint under feedback control in the walking stick insect? : I. Force measurements , 1981 .

[141]  J. Abbs,et al.  Compensatory responses to low magnitude loads applied to the lower lip during speech , 1981 .

[142]  C. Krarup Enhancement and diminution of mechanical tension evoked by staircase and by tetanus in rat muscle , 1981, The Journal of physiology.

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

[144]  R. Shaw,et al.  Psychophysics and ecometrics , 1981, Behavioral and Brain Sciences.

[145]  G. E. Goslow,et al.  Electrical activity and relative length changes of dog limb muscles as a function of speed and gait. , 1981, The Journal of experimental biology.

[146]  R. Stein,et al.  Nonlinear summation of contractions in cat muscles. I. Early depression , 1981, The Journal of general physiology.

[147]  A. Koivo,et al.  Control of robotic manipulator with adaptive controller , 1981, 1981 20th IEEE Conference on Decision and Control including the Symposium on Adaptive Processes.

[148]  S. Andreassen,et al.  Regulation of soleus muscle stiffness in premammillary cats: intrinsic and reflex components. , 1981, Journal of neurophysiology.

[149]  S. Andreassen,et al.  Limitations in the servo-regulation of soleus muscle stiffness in premammillary cats , 1981 .

[150]  Virginia A. Diggles,et al.  Control theories in motor behavior , 1982 .

[151]  J. W. Aldridge,et al.  Nonlinear properties of stretch reflex studied in the decerebrate cat. , 1982, Journal of neurophysiology.

[152]  W. D. Letbetter,et al.  Anatomy and innervation patterns of cat lateral gastrocnemius and plantaris muscles. , 1982, The American journal of anatomy.

[153]  W. D. Letbetter,et al.  A histochemical analysis of identified compartments of cat lateral gastrocnemius muscle , 1982, The Anatomical record.

[154]  V. V. Krishnan,et al.  Biological optimality: Comparison of generalized feasibility and optimality conditions for linear, error-minimizing systems , 1982 .

[155]  H. Cruse,et al.  The control system of the femur-tibia joint in the standing leg of a walking stick insect Carausius morosus , 1983 .