Phasic Modulation of Reflexes during Rhythmic Activity

  • Serge Rossignol
  • T. Drew
Part of the Wenner-Gren Center International Symposium Series book series (WGS)


Ten years ago (Forssberg et al, 1975, 1976, 1977) experiments in the chronic spinal kitten were designed to evaluate how reactions to perturbations of a limb during walking were integrated into the locomotor pattern. It was observed that, when the dorsum of the hindfoot contacted a rod during swing, the whole leg elegantly hyperflexed over the obstacle. However, when the same region of the foot was touched by the rod during stance, mimicking for instance a twig flying back, there was no response in the flexor muscles but instead an increase in the activity of the extensor muscles. To ensure that the very same stimulation did indeed produce two opposite responses, the experiments were repeated with electrical stimulation of the dorsum of the hindfoot. This stimulation also induced an ipsilateral flexion response only during the swing phase of the step cycle and an ipsilateral extension response during the stance phase. This phenomenon, in keeping with the past terminology, was referred to as reflex reversal. Implied in this expression was the fact that the transmission of a given pathway was effective in one phase and ineffective in the opposite phase of the movement. This applied equally to pathways exciting flexors, mainly during the swing phase, as well as to pathways exciting extensors, mainly in the stance phase, although the latencies of the responses elicited through these pathways could be slightly different.


Stance Phase Swing Phase Step Cycle Excitatory Response Reflex Pathway 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


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  1. Andersson, O., Forssberg, H., Grillner, S. and Lindquist, M, (1978). Phasic Gain Control of the Transmission in Cutaneous Reflex Pathways to Motoneurones during “Fictive” Locomotion. Brain Res., 149, 503–507.CrossRefPubMedGoogle Scholar
  2. Aoki, M. and McIntyre, A.K. (1975). Cortical and Long Spinal Actions on Lumbosacral Motoneurones in the Cat. J. Physiol. (Lond.), 251, 569–587.CrossRefGoogle Scholar
  3. Bayev, K.V. and Kostyuk, P.G. (1981). Primary Afferent Depolarization Evoked by the Activity of Spinal Scratching Generator. Neurosci., 6, 205–215.CrossRefGoogle Scholar
  4. Bayev, K.V. and Kostyuk, P.G. (1982). Polarization of Primary Afferent Terminals of Lumbosacral Cord Elicited by the Activity of Spinal Locomotor Generator. Neurosci., 7, 1401–1409.CrossRefGoogle Scholar
  5. Bélanger, M. and Patla, A.E. (1984). Corrective Responses to Perturbation Applied during Walking in Humans. Neurosci. Lett., 49, 291–295.CrossRefPubMedGoogle Scholar
  6. Cabelguen, J.-M., Orsal, D., Perret, C. and Zattara, M. (1981). Central Pattern Generation of Forelimb and Hindlimb Locomotor Activities in the Cat. In Regulatory Functions of the CNS. Motion and organization Principles. Advances in Physiological Sciences. Vol1. (eds. J. Szentagothai, M. Palkovits and J. Hamori.). Akademiai Kiado, Budapest, 199–211.CrossRefGoogle Scholar
  7. Crenna, P. and Frigo, C. (1984). Evidence of Phase-dependent Nociceptive Reflexes during Locomotion in Man. Exp. Neurol., 85, 336–345.CrossRefPubMedGoogle Scholar
  8. Drew, T. and Rossignol, S. (1984). Phase Dependent Responses Evoked in Limb Muscles by Stimulation of the Medullary Reticular Formation during Locomotion in Thalamic Cats. J. Neurophy siol., 52, 653–675.Google Scholar
  9. Drew, T. and Rossignol, S. (1985). Forelimb Responses to Cutaneous Nerve Stimulation during Locomotion in Intact Cats. Brain Res., 329, 323–328.CrossRefPubMedGoogle Scholar
  10. Duysens, J. (1977). Reflex Control of Locomotion as Revealed by Stimulation of Cutaneous Afferents in Spontaneously Walking Premammillary Cats. J. Neurophysiol., 40, 737–751.PubMedGoogle Scholar
  11. Duysens, J. and Loeb, G.E. (1980). Modulation of Ipsi- and Contralateral Reflex Responses in Unrestrained Walking Cats. J. Neurophysiol., 44, 1024–1037.PubMedGoogle Scholar
  12. Duysens, J., Loeb, G.E. and Weston, B.J. (1980). Crossed Flexor Reflex Responses and their Reversal in Freely Walking Cats. Brain Res., 197, 538–542.CrossRefPubMedGoogle Scholar
  13. Duysens, J. and Pearson, K.G. (1976). The Role of Cutaneous Afferents from the Distal Hind-limb in the Regulation of the Step Cycle of Thalamic Cats. Exp. Brain Res., 24, 245–255.CrossRefPubMedGoogle Scholar
  14. Duysens J., and Stein, R.B. (1978). Reflexes Induced by Nerve Stimulation in Walking Cats with Implanted Cuff Electrodes. Exp. Brain Res. 32, 213–224.CrossRefPubMedGoogle Scholar
  15. Forssberg, H. (1979). Stumbling Corrective Reaction: A Phase-dependent Compensatory Reaction during Locomotion. J. Neurophysiol., 42, 936–953.PubMedGoogle Scholar
  16. Forssberg, H. (1981). Phasic Gating of Cutaneous Reflexes during Locomotion. In Muscle Receptors and movement, (eds. A. Taylor and A. Prochazka). MacMillan, London, 403–412CrossRefGoogle Scholar
  17. Forssberg, H. (1985). Phase dependent step adaptations during human locomotion. In Feedback and Motor Control, (eds. W.J.P. Barnes and M.H. Gladden). Croom Helm Ltd., London (in press).Google Scholar
  18. Forssberg, H., Grillner, S. and Rossignol, S. (1975). Phase Dependent Reflex Reversal during Walking in Chronic Spinal Cats. Brain Res., 85, 103–107.CrossRefPubMedGoogle Scholar
  19. Forssberg, H., Grillner, S. and Rossignol S. (1977). Phasic Gain Control of Reflexes from the Oorsum of the Paw during Spinal Locomotion. Brain Res., 132, 121–139.CrossRefPubMedGoogle Scholar
  20. Forssberg, H., Grillner, S., Rossignol, S. and Wallén, P. (1976). Phasic Control of Reflexes during Locomotion in Vertebrates. In The Neural Control of Locomotion. (eds. R.M. Herman, S. Grillner, P. Stein and D. Stuart). Plenum Press, New York, 647–674.CrossRefGoogle Scholar
  21. Gauthier, L. and Rossignol, S. (1981). Contralateral Hindlimb Responses to Cutaneous Stimulation during Locomotion in High Decerebrate Cats. Brain Res., 207, 303–320.CrossRefPubMedGoogle Scholar
  22. Grillner, S. and Rossignol, S. (1978). Contralateral Reflex Reversal Controlled by Limb Position in the Acute Spinal Cat Injected with Clonidine i.v. Brain Res., 144, 411–414.CrossRefPubMedGoogle Scholar
  23. Grillner, S., Rossignol, S. and Wallén, P. (1977). The Adaptation of a Reflex Response to the Ongoing Phase of Locomotion in Fish. Exp. Brain Res., 30, 1–11.CrossRefPubMedGoogle Scholar
  24. Julien, C, Barbeau, H. and Rossignol, S. (1982). Gain Changes in Cutaneous Reflexes during Locomotion in the Adult Chronic Spinal Cat. Soc. Neurosci. Abstr., 8, 168.Google Scholar
  25. Kanda, K. and Sato, H (1983). Reflex Responses of Human Thigh Muscles to Non-noxious Sural Stimulation during Stepping. Brain Res., 288, 378–380.CrossRefPubMedGoogle Scholar
  26. Lisin, V.V., Frankstein, S.I. and Rechtmann, M.B. (1973). The Influence of Locomotion on Flexor Reflex of the Hindlimb in Cat and Man. Exp. Neurol., 38, 180–183.CrossRefPubMedGoogle Scholar
  27. Lloyd, D.P.C. and Mclntyre, A.K. (1948). Analysis of Forelimbhindlimb Reflex Activity in Acutely Decapitate Cats, J. Neuro physiol., 11, 455–470.Google Scholar
  28. Lund, J.P. and Olsson, K.A. (1983). The Importance of Reflexes and their Control during Jaw Movement. Trends in Neurosci., 6, 458–463.CrossRefGoogle Scholar
  29. Lund J.P. and Rossignol, S. (1981). Modulation of the Amplitude of the Digastric Jaw Opening Reflex during the Masticatory Cycle. Neuroscience, 6, 95–98.CrossRefPubMedGoogle Scholar
  30. Lund, J., Rossignol, S. and Murakami, T. (1981). Interactions betwen the Jaw Opening Reflex and Mastication. Can. J. Physiol. Pharmacol., 59, 683–690.CrossRefPubMedGoogle Scholar
  31. Matsukawa, K., Kamei, H., Minoda, K. and Udo, M. (1982). Inter-limb Coordination in Cat Locomotion Investigated with Perturbation. I. Behavioral and Electromyographic Study on Symmetric Limbs of Decerebrate and Awake Walking Cats. Exp. Brain Res., 46, 425–437.CrossRefPubMedGoogle Scholar
  32. Miller, S., Ruit, J.B. and van der Méché, F.G.A. (1977). Reversal of Sign of Long Spinal Reflexes Dependent on the Phase of the Step Cycle in the High Decerebrate Cat. Brain Res., 128, 447–459.CrossRefPubMedGoogle Scholar
  33. Nashner, L.M., Woollacott, M. and Tuma, G. (1979). Organization of Rapid Responses to Postural and Locomotor-Like Perturbations of Standing Man. Exp. Brain Res., 36, 463–476.CrossRefPubMedGoogle Scholar
  34. Perret, C. and Cabelguen, J.-M. (1980). Main Characteristics of the Hindlimb Locomotor Cycle in the Decorticate Cat with Special Reference to Bifunctional Muscles. Brain Res., 187, 333–352.CrossRefPubMedGoogle Scholar
  35. Rossignol, S. and Drew, T. (1985). Interactions of Segmental and Suprasegmental Inputs with the Spinal Pattern Generator of Locomotion. In Feedback and Motor Control. (eds. W.J.P. Barnes and M.H. Gladden). Croom Helm Ltd., London (in press).Google Scholar
  36. Rossignol, S. and Gauthier, L. (1980). An Analysis of Mechanisms Controlling the Reversal of Crossed Spinal Reflexes. Brain Res., 182, 31–45.CrossRefPubMedGoogle Scholar
  37. Rossignol, S., Julien, C. and Gauthier, L. (1981a). Stimulus-response Relationships during Locomotion. Can. J. Physiol., 59, 667–674.CrossRefGoogle Scholar
  38. Rossignol, S., Gauthier, L., Julien, C. and Lund, J.P. (1981b). State Dependent Responses during Locomotion. In Muscle Receptors and Movement. (eds. A. Taylor, A. Prochazka). MacMillan Publis., 389–402.CrossRefGoogle Scholar
  39. Schomburg, E.D. and Behrends, H.B. (1978). Phasic Control of the Transmission in the Excitatory and Inhibitory Reflex Pathways from Cutaneous Afferents to Motoneurones during Fictive Locomotion in Cats. Neurosci. Lett., 8, 277–282.CrossRefPubMedGoogle Scholar
  40. Schomburg, E.D., Behrends, H.B. and Steffens, H. (1981). Changes in Segmental and Propriospinal Reflex Pathways during Spinal Locomotion. In Muscle Receptors and Movements. (eds. A. Taylor and A. Prochazka). MacMillan, London, 413–425.CrossRefGoogle Scholar
  41. Schomburg, E.D., Roesler, J. and Meinck, H.M. (1977). Phase-dependent Transmission in the Excitatory Propriospinal Reflex Pathway from Forelirab Afferents to Lumbar Motoneurones during Fictive Locomotion. Neurosci. Lett., 4, 249–252.CrossRefPubMedGoogle Scholar
  42. Sherrington, C.S. (1898). Decerebrate Rigidity and Reflex Coordination of Movements. J. Physiol., 22, 319–332.PubMedCentralCrossRefPubMedGoogle Scholar
  43. Wallén, P. (1980). On the Mechanisms of a Phase Dependent Reflex Occurring during Locomotion in Dogfish. Exp. Brain Res. 39, 193–202.CrossRefPubMedGoogle Scholar
  44. Wand P., Prochazka, A., and Sontag, K.H. (1980). Neuromuscular Responses to Gait Perturbations in Freely Moving Cats. Exp. Brain Res., 38, 109–114.CrossRefPubMedGoogle Scholar

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© The Wenner-Gren Center 1986

Authors and Affiliations

  • Serge Rossignol
  • T. Drew

There are no affiliations available

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