Neural Darwinism in the Mammalian Spinal Cord

A conceptual framework around which the spinal control of locomotion is viewed and often studied has focused on the property of “Central Pattern Generation”, i.e. the generation of oscillating efferent patterns from the lumbosacral spinal cord in the absence of either supraspinal or sensory oscillating input (Grillner, 1973; 1982; Severston, 1999). An alternative, or at least complementary, concept that may help to understand spinally-controlled locomotion has been presented by Edelman and colleagues (Edelman, 1978; Edelman, 1981; Edelman and Reeke, 1982; Edelman and Finkel, 1984) to account for adaptive events in the brain. They theorized that “the brain is dynamically organized into cellular populations containing individually varied networks, the structure and function of which are selected by different means during development and behavior”. The cellular populations are proposed to be collections of hundreds to thousands of strongly interconnected neurons acting as functional units. The key elements of this Neuronal Group Selection concept, often referred to as Neural Darwinism, are a) “Reentrant” structure and function involving a continuous spatial temporal representation of sensory information and a mechanism for continuous updating of the selection of neuronal groups that can generate a given motor synergy or pattern and, b) “Degeneracy” indicating that there must be variance in the neuronal groups that generate a movement pattern if learning and selective matching of sensory and motor processes are to occur. This concept, therefore, also assumes that different neuronal groups can accomplish the same function.

[1]  Konstantin V. Baev,et al.  Biological Neural Networks: Hierarchical Concept of Brain Function , 1998, Birkhäuser Boston.

[2]  S. Rossignol,et al.  Recovery of locomotion after chronic spinalization in the adult cat , 1987, Brain Research.

[3]  V. Edgerton,et al.  Neural influence on slow muscle properties: Inactivity with and without cross‐reinnervation , 1996, Muscle & nerve.

[4]  V R Edgerton,et al.  Retention of hindlimb stepping ability in adult spinal cats after the cessation of step training. , 1999, Journal of neurophysiology.

[5]  S. S. Tower,et al.  Function and structure in the chronically isolated lumbo‐sacral spinal cord of the dog , 1937 .

[6]  V R Edgerton,et al.  A physiological basis for the development of rehabilitative strategies for spinally injured patients. , 1991, The Journal of the American Paraplegia Society.

[7]  V R Edgerton,et al.  Chronic spinal cord-injured cats: surgical procedures and management. , 1992, Laboratory animal science.

[8]  S. Rossignol,et al.  Initiation and modulation of the locomotor pattern in the adult chronic spinal cat by noradrenergic, serotonergic and dopaminergic drugs , 1991, Brain Research.

[9]  S. Grillner,et al.  Neuronal Control of LocomotionFrom Mollusc to Man , 1999 .

[10]  G M Edelman,et al.  Selective networks capable of representative transformations, limited generalizations, and associative memory. , 1982, Proceedings of the National Academy of Sciences of the United States of America.

[11]  C. Bouchard,et al.  Physical Activity, Fitness, and Health: International Proceedings and Consensus Statement , 1994 .

[12]  J. Duysens,et al.  Load-regulating mechanisms in gait and posture: comparative aspects. , 2000, Physiological reviews.

[13]  A Selverston,et al.  General principles of rhythmic motor pattern generation derived from invertebrate CPGs. , 1999, Progress in brain research.

[14]  V R Edgerton,et al.  Locomotor capacity attributable to step training versus spontaneous recovery after spinalization in adult cats. , 1998, Journal of neurophysiology.

[15]  G. Edelman Neural Darwinism: The Theory Of Neuronal Group Selection , 1989 .

[16]  J M Macpherson,et al.  Weight support and balance during perturbed stance in the chronic spinal cat. , 1999, Journal of neurophysiology.

[17]  Serge Rossignol,et al.  Effects of Intrathecal α1- and α2-Noradrenergic Agonists and Norepinephrine on Locomotion in Chronic Spinal Cats , 1998 .

[18]  G. Edelman Group selection and phasic reentrant signaling a theory of higher brain function , 1982 .

[19]  V R Edgerton,et al.  Training effects on soleus of cats spinal cord transected (T12–13) as adults , 1998, Muscle & nerve.

[20]  V. Reggie Edgerton,et al.  Coordination of motor pools controlling the ankle musculature in adult spinal cats during treadmill walking , 1991, Brain Research.

[21]  V. Dietz,et al.  Locomotor capacity of spinal cord in paraplegic patients , 1995, Annals of neurology.

[22]  Sholl Da Organization of the Cerebral Cortex , 1967 .

[23]  P. Stein Chapter 23 Central Pattern Generators and Interphyletic Awareness , 1999 .

[24]  V. Edgerton,et al.  Mechanical and morphological properties of chronically inactive cat tibialis anterior motor units. , 1991, The Journal of physiology.

[25]  K. Pearson Proprioceptive regulation of locomotion , 1995, Current Opinion in Neurobiology.

[26]  V R Edgerton,et al.  Potential of adult mammalian lumbosacral spinal cord to execute and acquire improved locomotion in the absence of supraspinal input. , 1992, Journal of neurotrauma.

[27]  M. Gorassini,et al.  Models of ensemble firing of muscle spindle afferents recorded during normal locomotion in cats , 1998, The Journal of physiology.

[28]  R. J. Gregor,et al.  Weight-bearing hindlimb stepping in treadmill-exercised adult spinal cats , 1990, Brain Research.

[29]  V R Edgerton,et al.  Hindlimb locomotor and postural training modulates glycinergic inhibition in the spinal cord of the adult spinal cat. , 1999, Journal of neurophysiology.

[30]  Kathleen E. Norman,et al.  Review : Walking After Spinal Cord Injury: Control and Recovery , 1998 .

[31]  L. Rowell,et al.  Exercise : regulation and integration of multiple systems , 1996 .

[32]  A Prochazka,et al.  Ensemble firing of muscle afferents recorded during normal locomotion in cats , 1998, The Journal of physiology.

[33]  V. Reggie Edgerton,et al.  Does Motor Learning Occur in the Spinal Cord? , 1997 .

[34]  H. Barbeau,et al.  Modulation of Locomotor Patterns and Spasticity with Clonidine in Spinal Cord Injured Patients , 1991, Canadian Journal of Neurological Sciences / Journal Canadien des Sciences Neurologiques.

[35]  B. Dobkin,et al.  Human lumbosacral spinal cord interprets loading during stepping. , 1997, Journal of neurophysiology.

[36]  S. Grillner Control of Locomotion in Bipeds, Tetrapods, and Fish , 1981 .

[37]  R. Kretchmar Exercise and Sport Science , 1989 .

[38]  K. Pearson,et al.  Control of Posture and Locomotion , 1973, Advances in Behavioral Biology.

[39]  Gerald M. Edelman,et al.  Dynamic aspects of neocortical function , 1984 .

[40]  S. Rossignol,et al.  The effects of serotonergic drugs on the locomotor pattern and on cutaneous reflexes of the adult chronic spinal cat , 1990, Brain Research.

[41]  J M Macpherson,et al.  Stance control in the chronic spinal cat. , 1994, Journal of neurophysiology.

[42]  S. Rossignol,et al.  Phasic gain control of reflexes from the dorsum of the paw during spinal locomotion , 1977, Brain Research.

[43]  C. Capaday,et al.  Difference in the amplitude of the human soleus H reflex during walking and running. , 1987, The Journal of physiology.

[44]  V R Edgerton,et al.  Full weight-bearing hindlimb standing following stand training in the adult spinal cat. , 1998, Journal of neurophysiology.

[45]  E. Marder,et al.  Switching neurons are integral members of multiple oscillatory networks , 1994, Current Biology.

[46]  S. Rossignol,et al.  The effects of clonidine and yohimbine on locomotion and cutaneous reflexes in the adult chronic spinal cat , 1987, Brain Research.

[47]  S. Grillner,et al.  Neuronal Control of Locomotion 'From Mollusc to Man ' , 1999 .

[48]  V R Edgerton,et al.  Use-dependent plasticity in spinal stepping and standing. , 1997, Advances in neurology.

[49]  R. J. Gregor,et al.  Effects of training on the recovery of full-weight-bearing stepping in the adult spinal cat , 1986, Experimental Neurology.

[50]  S. Rossignol,et al.  Pharmacology of locomotion: an account of studies in spinal cats and spinal cord injured subjects. , 1993, The Journal of the American Paraplegia Society.

[51]  A. Wernig,et al.  Laufband Therapy Based on‘Rules of Spinal Locomotion’is Effective in Spinal Cord Injured Persons , 1995, The European journal of neuroscience.

[52]  S. Grillner Locomotion in the Spinal Cat , 1973 .

[53]  V R Edgerton,et al.  The plasticity of skeletal muscle: effects of neuromuscular activity. , 1991, Exercise and sport sciences reviews.

[54]  S. Rossignol,et al.  Early locomotor training with clonidine in spinal cats. , 1998, Journal of neurophysiology.

[55]  S. Rossignol,et al.  An analysis of mechanisms controlling the reversal of crossed spinal reflexes , 1980, Brain Research.

[56]  G. Edelman,et al.  The Mindful Brain: Cortical Organization and the Group-Selective Theory of Higher Brain Function , 1978 .

[57]  B. Dobkin,et al.  Can the mammalian lumbar spinal cord learn a motor task? , 1994, Medicine and science in sports and exercise.