Abstract
During locomotion sensory information from cutaneous and muscle receptors is continuously integrated with the locomotor central pattern generator (CPG) to generate an appropriate motor output to meet the demands of the environment. Sensory signals from peripheral receptors can strongly impact the timing and amplitude of locomotor activity. This sensory information is gated centrally depending on the state of the system (i.e., rest vs. locomotion) but is also modulated according to the phase of a given task. Consequently, if one is to devise biologically relevant walking models it is imperative that these sensorimotor interactions at the spinal level be incorporated into the control system.
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Abelew TA, Miller MD, Cope TC, Nichols TR (2000) Local loss of proprioception results in disruption of interjoint coordination during locomotion in the cat. J Neurophysiol 84:2709–2714
Abraham LD, Marks WB, Loeb GE (1985) The distal hindlimb musculature of the cat. Cutaneous reflexes during locomotion. Exp Brain Res 58:594–603
Andersen JB, Sinkjaer T (1999) The stretch reflex and H-reflex of the human soleus muscle during walking. Motor Control 3:151–157
Anderson JH (1974) Dynamic characteristics of Golgi tendon organs. Brain Res 67:531–537
Andersson O, Grillner S (1981) Peripheral control of the cat’s step cycle. I. Phase dependent effects of ramp-movements of the hip during “fictive locomotion” Acta Physiol Scand 113:89–101
Andersson O, Grillner S (1983) Peripheral control of the cat’s step cycle. II. Entrainment of the central pattern generators for locomotion by sinusoidal hip movements during “fictive locomotion” Acta Physiol Scand 118:229–239
Angel MJ, Guertin P, Jimenez T, McCrea DA (1996) Group I extensor afferents evoke disynaptic EPSPs in cat hindlimb extensor motorneurones during fictive locomotion. J Physiol 494:851–861
Barbeau H, Rossignol S (1987) Recovery of locomotion after chronic spinalization in the adult cat. Brain Res 412:84–95
Bélanger M, Drew T, Rossignol S (1988) Spinal locomotion: a comparison of the kinematics and the electromyographic activity in the same animal before and after spinalization. Acta Biol Hungarica 39:151–154
Bélanger M, Drew T, Provencher J, Rossignol S (1996) A comparison of treadmill locomotion in adult cats before and after spinal transection. J Neurophysiol 76:471–491
Bem T, Cabelguen JM, Ekeberg O, Grillner S (2003) From swimming to walking: a single basic network for two different behaviors. Biol Cybern 88:79–90
Bennett DJ, De Serres SJ, Stein RB (1996) Gain of the triceps surae stretch reflex in decerebrate and spinal cats during postural and locomotor activities. J Physiol 496:837–850
Bessou P, Dupui P, Cabelguen JM, Joffroy M, Montoya R, Pages B (1989) Discharge patterns of gamma motoneurone populations of extensor and flexor hindlimb muscles during walking in the thalamic cat. Prog Brain Res 80:37–45
Bizzi E, Giszter SF, Loeb E, Mussa-Ivaldi FA, Saltiel P (1995) Modular organization of motor behavior in the frog’s spinal cord. Trends Neurosci 18:442–446
Booth V, Rinzel J, Kiehn O (1997) Compartmental model of vertebrate motoneurons for Ca2+-dependent spiking and plateau potentials under pharmacological treatment. J Neurophysiol 78:3371–3385
Bouyer LJG, Rossignol S (2003a) Contribution of cutaneous inputs from the hindpaw to the control of locomotion: 1. Intact cats. J Neurophysiol 90:3625–3639
Bouyer LJG, Rossignol S (2003b) Contribution of cutaneous inputs from the hindpaw to the control of locomotion: 2. Spinal cats. J Neurophysiol 90:3640–3653
Brown TG (1914) On the nature of the fundamental activity of the nervous centres together with an analysis of the conditioning of rhythmic activity in progression and a theory of the evolution of function in the nervous system. J Physiol 48: 18–46
Brown IE, Loeb GE (2000) Measured and modeled properties of mammalian skeletal muscle: IV. dynamics of activation and deactivation. J Muscle Res Cell Motil 21:33–47
Brown IE, Scott SH, Loeb GE (1996) Mechanics of feline soleus: II. Design and validation of a mathematical model. J Muscle Res Cell Motil 17:221–233
Brown IE, Cheng EJ, Loeb GE (1999) Measured and modeled properties of mammalian skeletal muscle. II. The effects of stimulus frequency on force-length and force-velocity relationships. J Muscle Res Cell Motil 20:627–643
Brownstone RM, Jordan LM, Kriellaars DJ, Noga BR, Shefchyk SJ (1992) On the regulation of repetitive firing in lumbar motoneurones during fictive locomotion in the cat. Exp Brain Res 90:441–455
Buford JA, Smith JL (1990) Adaptive control for backward quadrupedal walking. II.Hindlimb muscle synergies. J Neurophysiol 64:756–766
Buford JA, Smith JL (1993) Adaptive control for backward quadrupedal walking: III. Stumbling corrective reactions and cutaneous reflex sensitivity. J Neurophysiol 70: 1102–1114
Burke RE (1999) The use of state-dependent modulation of spinal reflexes as a tool to investigate the organization of spinal interneurons. Exp Brain Res 128:263–277
Burke RE, Degtyarenko AM, Simon ES (2001) Patterns of locomotor drive to motoneurons and last-order interneurons: clues to the structure of the CPG. J Neurophysiol 86: 447–462
Butera Jr RJ, Rinzel J, Smith JC (1999) Models of respiratory rhythm generation in the pre-Botzinger complex. II. Populations Of coupled pacemaker neurons. J Neurophysiol 82:398–415
Cabelguen J-M, Orsal D, Perret C, Zattara M (1981) Central pattern generation of forelimb and hindlimb locomotor activities in the cat. In: Szentagothai J, Palkovits M, Hamori J (eds) Regulatory functions of the CNS, principles of motion and organization. Adv Physiol Sci 1: Akadamiai Kiado, Budapest, pp 199–211
Carrier L, Brustein L, Rossignol S (1997) Locomotion of the hindlimbs after neurectomy of ankle flexors in intact and spinal cats: model for the study of locomotor plasticity. J Neurophysiol 77:1979–1993
Chen WJ, Poppele RE (1978) Small-signal analysis of response of mammalian muscle spindles with fusimotor stimulation and a comparison with large-signal responses. J Neurophysiol 41:15–27
Conway BA, Hultborn H, Kiehn O (1987) Proprioceptive input resets central locomotor rhythm in the spinal cat. Exp Brain Res 68:643–656
Cruse H (1990) What mechanisms coordinate leg movement in walking arthropods? Trends Neurosci 13:15–21
Cruse H, Bartling C, Cymbalyuk G, Dean J, Dreifert M (1995) A modular artificial neural net for controlling a six-legged walking system. Biol Cybern 72:421–430
Cruse H, Dean J, Kindermann T, Schmitz J, Schumm M (1998) Simulation of complex movements using artificial neural networks. Z Naturforsch [C ] 53:628–638
Degtyarenko AM, Simon ES, Burke RE (1996) Differential modulation of disynaptic cutaneous inhibition and excitation in ankle flexor motoneurons during fictive locomotion. J Neurophysiol 76:2972–2985
Dietz V, Colombo G (1998) Influence of body load on the gait pattern in Parkinson’s disease. Mov Disorders 13:255–261
Dietz V, Duysens J (2000) Significance of load receptor input during locomotion: a review. Gait Posture 11:102–110
Donelan JM, Pearson KG (2004a) Contribution of force feedback to ankle extensor activity in decerebrate walking cats. J Neurophysiol 92:2093–2104
Donelan JM, Pearson KG (2004b) Contribution of sensory feedback to ongoing ankle extensor activity during the stance phase of walking. Can J Physiol Pharmacol 82:589–598
Drew T (1988) Motor cortical cell discharge during voluntary gait modification. Brain Res 457:181–187
Drew T, Rossignol S (1987) A kinematic and electromyographic study of cutaneous reflexes evoked from the forelimb of unrestrained walking cats. J Neurophysiol 57:1160–1184
Drew T, Jiang W, Kably B, Lavoie S (1996) Role of the motor cortex in the control of visually triggered gait modifications. Can J Physiol Pharmacol 74:426–442
Durbaba R, Taylor A, Rawlinson SR, Ellaway PH (2003) Static fusimotor action during locomotion in the decerebrated cat revealed by cross-correlation of spindle afferent activity. Exp Physiol 88:285–296
Duysens J, Loeb GE (1980) Modulation of ipsi-and contralateral reflex responses in unrestrained walking cats. J Neurophysiol 44:1024–1037
Duysens J, Pearson KG (1980) Inhibition of flexor burst generation by loading ankle extensor muscles in walking cats. Brain Res 187:321–332
Duysens J, Tax AAM, Trippel M, Dietz V (1992) Phase-dependent reversal of reflexly induced movements during human gait. Exp Brain Res 90:404–414
Duysens J, Clarac F, Cruse H (2000) Load-regulating mechanisms in gait and posture: comparative aspects. Physiol Rev 80:83–133
Ekeberg O (1993) A combined neuronal and mechanical model of fish swimming. Biol Cybern 69:363–374
Ekeberg O, Grillner S (1999) Simulations of neuromuscular control in lamprey swimming. Philos Trans R Soc Lond B Biol Sci 354:895–902
Ekeberg O, Pearson K (2005) Computer simulation of stepping in the hind legs of the cat: an examination of mechanisms regulating the stance-to-swing transition. J Neurophysiol 94:4256–4268
Ekeberg O, Grillner S, Lansner A (1995) The neural control of fish swimming studied through numerical simulations. Adapt Behav 3:363–384
Ekeberg O, Blumel M, Buschges A (2004) Dynamic simulation of insect walking. Arthropod Struct Develop 33:287–300
Engberg I, Lundberg A (1969) An electromyographic analysis of muscular activity in the hindlimb of the cat during unrestrained locomotion. Acta Physiol Scand 75:614–630
Fleshman JW, Lev-Tov A, Burke RE (1984) Peripheral and central control of flexor digitorium longus and flexor hallucis longus motoneurons: the synaptic basis of functional diversity. Exp Brain Res 54:133–149
Forssberg H (1979) Stumbling corrective reaction: a phase-dependent compensatory reaction during locomotion. J Neurophysiol 42:936–953
Forssberg H, Grillner S, Rossignol S (1975) Phase dependent reflex reversal during walking in chronic spinal cats. Brain Res 85:103–107
Forssberg H, Grillner S, Rossignol S, Wallen P (1976) Phasic control of reflexes during locomotion in vertebrates. In: Herman RM, Grillner S, Stein PSG, Stuart DG (eds) Neural control of locomotion. Plenum, New York, pp 647–674
Forssberg H, Grillner S, Rossignol S (1977) Phasic gain control of reflexes from the dorsum of the paw during spinal locomotion. Brain Res 132:121–139
Forssberg H, Grillner S, Halbertsma J (1980a) The locomotion of the low spinal cat. I. Coordination within a hindlimb. Acta Physiol Scand 108:269–281
Forssberg H, Grillner S, Halbertsma J, Rossignol S (1980b) The locomotion of the low spinal cat: II. Interlimb coordination. Acta Physiol Scand 108:283–295
Giuliani CA, Smith JL (1987) Stepping behaviors in chronic spinal cats with one hindlimb deafferented. J Neurosci 7:2537–2546
Goldberger ME (1977) Locomotor recovery after unilateral hindlimb deafferentation in cats. Brain Res 123:59–74
Goldberger ME (1983) Recovery of accurate limb movements after deafferentation in cats. In: Kao CC, Bunge RP, Reier PJ (eds) Spinal cord reconstruction. Raven, New York, pp 455–463
Goldberger ME (1987) Recovery of locomotion and postural reflexes after spinal cord deafferentation. Neuroscience 22:S6
Gorassini MA, Prochazka A, Hiebert GW, Gauthier MJA (1994) Corrective responses to loss of ground support during walking I. Intact cats. J Neurophysiol 71:603–609
Gossard J-P, Hultborn H (1991) The organization of the spinal rhythm generation in locomotion. In: Wernig A (ed) Restorative neurology. Elsevier, Amsterdam, pp 385–404
Gossard J-P, Brownstone RM, Barajon I, Hultborn H (1994) Transmission in a locomotor-related group Ib pathway from hindlimb extensor muscles in the cat. Exp Brain Res 98:213–228
Granat MH, Heller BW, Nicol DJ, Baxendale RH, Andrews BJ (1993) Improving limb flexion in FES gait using the flexion withdrawal response for the spinal cord injured person. J Biomed Eng 15:51–56
Gregory JE, Proske U (1979) The responses of Golgi tendon organs to stimulation of different combinations of motor units. J Physiol 295:251–262
Gregory JE, Proske U (1981) Motor unit contractions initiating impulses in a tendon organ in the cat. J Physiol 313:251–262
Gregory JE, Morgan DL, Proske U (1985) Site of impulse initiation in tendon organs of cat soleus muscle. J Neurophysiol 54:1383–1395
Grillner S (1981) Control of locomotion in bipeds, tetrapods, and fish. In: Brookhart JM, Mountcastle VB (eds) Handbook of physiology. The nervous system II. American Physiological Society, Bethesda, pp 1179–1236
Grillner S, Rossignol S (1978) On the initiation of the swing phase of locomotion in chronic spinal cats. Brain Res 146:269–277
Grillner S, Zangger P (1979) On the central generation of locomotion in the low spinal cat. Exp Brain Res 34:241–261
Grillner S, Zangger P (1984) The effect of dorsal root transection on the efferent motor pattern in the cat’s hindlimb during locomotion. Acta Physiol Scand 120:393–405
Guertin P, Angel MJ, Perreault M-C, McCrea DA (1995) Ankle extensor group I afferents excite extensors throughout the hindlimb during fictive locomotion in the cat. J Physiol 487:197–209
Halbertsma JM (1983) The stride cycle of the cat: the modelling of locomotion by computerized analysis of automatic recordings. Acta Physiol Scand Suppl. 521:1–75
Harkema SJ, Hurlay SL, Patel UK, Requejo PS, Dobkin BH, Edgerton VR (1997) Human lumbosacral spinal cord interpret loading during stepping. J Neurophysiol 77:797–811
Hasan Z (1983) A model of spindle afferent response to muscle stretch. J Neurophysiol 49:989–1006
Hiebert GW, Whelan PJ, Prochazka A, Pearson KG (1996) Contribution of hind limb flexor muscle afferents to the timing of phase transitions in the cat step cycle. J Neurophysiol 75:1126–1137
Hill AV (1938) The heat of shortening and the dynamic constants of muscle. Proc R Soc Lond B 126:136–195
Houk J, Simon W (1967) Responses of Golgi tendon organs to forces applied to muscle tendon. J Neurophysiol 30:1466-1481
Houk JC, Rymer WZ, Crago PE (1981) Dependence of dynamic response of spindle receptors on muscle length and velocity. J Neurophysiol 46:143–166
Hultborn H (2006) Spinal reflexes, mechanisms and concepts: from Eccles to Lundberg and beyond. Prog Neurobiol 78:215–232
Hunt CC (1990) Mammalian muscle spindle: peripheral mechanisms. Physiol Rev 70:643–663
Ijspeert AJ (2001) A connectionist central pattern generator for the aquatic and terrestrial gaits of a simulated salamander. Biol Cybern 84:331–348
Ijspeert AJ, Crespi A, Cabelguen JM (2005) Simulation and studies of salamander locomotion: applying neurobiological principles to the control of locomotion in robots. Neuroinformatics 3:171–195
Ivanenko YP, Poppele RE, Lacquaniti F (2006) Spinal cord maps of spatiotemporal alpha-motoneuron activation in humans walking at different speeds. J Neurophysiol 95:602–618
Ivashko DG, Prilutsky BI, Markin SN, Chapin JK, Rybak IA (2003) Modeling the spinal cord neural circuitry controlling cat hindlimb movement during locomotion. Neurocomputing 52-:621-29
Jami L (1992) Golgi tendon organs in mammalian skeletal muscle: functional properties and central actions. Physiol Rev 72:623–666
Jankowska E, Jukes MGM, Lund S, Lundberg A (1967) The effects of DOPA on the spinal cord. 6. Half centre organization of interneurones transmitting effects from the flexor reflex afferents. Acta Physiol Scand 70:389–402
Jayne BC (1988) Muscular mechanisms of snake locomotion: an electromyographic study of lateral undulation of the Florida banded water snake (Nerodia fasciata) and the yellow rat snake (Elaphe obsoleta). J Morphol 197:159–181
Kendall FP, McCreary EK, Provance PG (1993) Muscles: testing and function. Lippincot Williams& Wilkins, Baltimore
Kiehn O (2006) Locomotor circuits in the Mammalian spinal cord. Annu Rev Neurosci 29:279–306
Koshland GF, Smith JL (1989) Mutable and immutable features of paw-shake responses after hindlimb deafferentation in the cat. J Neurophysiol 62:162–173
Kriellaars DJ, Brownstone RM, Noga BR, Jordan LM (1994) Mechanical entrainment of fictive locomotion in the decerebrate cat. J Neurophysiol 71:1–13
Kuo PD, Eliasmith C (2005) Integrating behavioral and neural data in a model of zebrafish network interaction. Biol Cybern 93:178–187
Kuo AD, Donelan JM, Ruina A (2005) Energetic consequences of walking like an inverted pendulum: step-to-step transitions. Exerc Sport Sci Rev 33:88–97
LaBella L, Niechaj A, Rossignol S (1992) Low-threshold, short-latency cutaneous reflexes during fictive locomotion in the “semi-chronic” spinal cat. Exp Brain Res 91:236–248
Lafreniere-Roula M, McCrea DA (2005) Deletions of rhythmic motoneuron activity during fictive locomotion and scratch provide clues to the organization of the mammalian central pattern generator. J Neurophysiol 94:1120–1132
Lam T, Pearson KG (2001) Proprioceptive modulation of hip flexor activity during the swing phase of locomotion in decerebrate cats. J Neurophysiol 86:1321–1332
Lam T, Pearson KG (2002) The role of proprioceptive feedback in the regulation and adaptation of locomotor activity. Adv Exp Med Biol 508:343–355
Langlet C, Leblond H, Rossignol S (2005) The mid-lumbar segments are needed for the expression of locomotion in chronic spinal cats. J Neurophysiol 93:2474–2488
Lundberg A (1981) Half-centres revisited. In: Szentagothai J, Palkovits M, Hamori J (eds) Regulatory Functions of the CNS. Principles of Motion and Organization. Advances in Physiological Science, vol 1 Pergamon, Budapest, pp 155-67
Matthews PBC, Stein RB (1969) The sensitivity of muscle spindle afferents to small sinusoidal changes of length. J Physiol 200:723–744
McCrea DA, Shefchyk SJ, Stephens MJ, Pearson KG (1995) Disynaptic group: I. excitation of ankle extensor motoneurones during fictive locomotion in the cat. J Physiol 487:527–539
McFadyen BJ, Winter DA, Allard F (1994) Simulated control of unilateral, anticipatory locomotor adjustments during obstructed gait. Biol Cybern 72:151–160
McMahon TA (1984) A Lumped, Linear Model of the Spindle. Muscles, Reflexes, and Locomotion. Princeton University Press, Princeton , pp 152–154
Mileusnic MP, Loeb GE (2006) Mathematical models of proprioceptors. II. Structure and function of the Golgi tendon organ. J Neurophysiol 96:1789–1802
Mileusnic MP, Brown IE, Lan N, Loeb GE (2006) Mathematical models of proprioceptors. I. Control and transduction in the muscle spindle. J Neurophysiol 96:1772–1788
Moschovakis AK, Sholomenko GN, Burke RE (1991) Differential control of short latency cutaneous excitation in cat FDL motoneurons during fictive locomotion. Exp Brain Res 83:489–501
Moschovakis AK, Solodkin M, Burke RE (1992) Anatomical and physiological study of interneurons in an oligosynaptic cutaneous reflex pathway in the cat hindlimb. Brain Res 586:311–318
Mott FW, Sherrington CS (1895) Experiments upon the influence of sensory nerves upon movement and nutrition of the limbs: preliminary communication. Proc R Soc Lond B481-89
Ogihara N, Yamazaki N (2001) Generation of human bipedal locomotion by a bio-mimetic neuro–musculo-skeletal model. Biol Cybern 84:1–11
Orlovsky GN, Deliagina TG, Grillner S (1999) Neuronal control of locomotion: from mollusc to man. Oxford University Press, New York , pp 1–322
Pandy MG (2001) Computer modeling and simulation of human movement. Annu Rev Biomed Eng 3:245–273
Pandy MG (2003) Simple and complex models for studying muscle function in walking. Philos Trans R Soc Lond B Biol Sci 358:1501–1509
Paul C, Bellotti M, Jezernik S, Curt A (2005) Development of a human neuro-musculo-skeletal model for investigation of spinal cord injury. Biol Cybern 93:153–170
Pearson KG (2000) Neural adaptation in the generation of rhythmic behavior. Annu Rev Physiol 62:723–753
Pearson KG, Rossignol S (1991) Fictive motor patterns in chronic spinal cats. J Neurophysiol 66:1874–1887
Pearson KG, Misiaszek JE, Hulliger M (2003) Chemical ablation of sensory afferents in the walking system of the cat abolishes the capacity for functional recovery after peripheral nerve lesions. Exp Brain Res 150:50–60
Pearson K, Ekeberg O, Buschges A (2006) Assessing sensory function in locomotor systems using neuro-mechanical simulations. Trends Neurosci 29:625–631
Perret C, Cabelguen J-M (1976) Central and reflex participation in the timing of locomotor activations of a bifunctional muscle, the semi-tendinosus, in the cat. Brain Res 106:390–395
Perret C, 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
Pratt CA, Chanaud CM, Loeb GE (1991) Functionally complex muscles of the cat hindlimb. IV. Intramuscular distribution of movement command signals and cutaneous reflexes in broad, bifunctional thigh muscles. Exp Brain Res 85: 281–299
Prochazka A (1989) Sensorimotor gain control: a basic strategy of motor systems? Prog Neurobiol 33:281–307
Prochazka A (1993) Comparison of natural and artificial control of movement. IEEE Trans Rehab Eng 1:7–17
Prochazka A (1996) Proprioceptive feedback and movement regulation. In: Rowell LB, Sheperd JT (eds) Handbook of physiology. Sect 12. Exercise: regulation and integration of multiple systems. American Physiological Society, New York, pp 89–127
Prochazka A, Gorassini M (1998a) Ensemble firing of muscle afferents recorded during normal locomotion in cats. J Physiol 507(Pt 1):293–304
Prochazka A, Gorassini M (1998b) Models of ensemble firing of muscle spindle afferents recorded during normal locomotion in cats. J Physiol 507(Pt 1):277–291
Prochazka A, Hulliger M, Zangger P, Appenteng K (1985) “Fusimotor set” new evidence for alpha-independent control of gamma-motoneurones during movement in the awake cat. Brain Res 339:136–140
Prochazka A, Gritsenko V, Yakovenko S (2002) Sensory control of locomotion: reflexes versus higher-level control. Adv Exp Med Biol 508:357–367
Quevedo J, Fedirchuk B, Gosgnach S, McCrea DA (2000) Group I disynaptic excitation of cat hindlimb flexor and bifunctional motoneurones during fictive locomotion. J Physiol 525:549–564
Quevedo J, Stecina K, Gosgnach S, McCrea DA (2005a) Stumbling corrective reaction during fictive locomotion in the cat. J Neurophysiol 94:2045–2052
Quevedo J, Stecina K, McCrea DA (2005b) Intracellular analysis of reflex pathways underlying the stumbling corrective reaction during fictive locomotion in the cat. J Neurophysiol 94:2053–2062
Rossignol S (1996) Neural control of stereotypic limb movements. In: Rowell LB, Sheperd JT (eds) Handbook of physiology, Sect. 12. Exercise: Regulation and Integration of Multiple Systems. Oxford University Press, New York, pp 173–216
Rossignol S, Saltiel P, Perreault M-C, Drew T, Pearson KG, Bélanger M (1993) Intralimb and interlimb coordination in the cat during real and fictive rhythmic motor programs. Neurosciences 5:67–75
Rossignol S, Bélanger M, Chau C, Giroux N, Brustein E, Bouyer L, Grenier C-A, Drew T, Barbeau H, Reader T (2000) The spinal cat. In: Kalb RG, Strittmatter SM (eds) Neurobiology of spinal cord injury. Humana, Totowa, pp 57–87
Rossignol S, Dubuc R, Gossard JP (2006) Dynamic sensorimotor interactions in locomotion. Physiol Rev 86:89–154
Rybak IA, Shevtsova NA, St John WM, Paton JF, Pierrefiche O (2003) Endogenous rhythm generation in the pre-Botzinger complex and ionic currents: modelling and in vitro studies. Eur J Neurosci 18:239–257
Rybak IA, Shevtsova NA, Lafreniere-Roula M, McCrea DA (2006a) Modelling spinal circuitry involved in locomotor pattern generation: insights from deletions during fictive locomotion. J Physiol doi: 10.1113/jphysiol.2006.118703 (in press)
Rybak IA, Stecina K, Shevtsova NA, McCrea DA (2006b) Modelling spinal circuitry involved in locomotor pattern generation: insights from the effects of afferent stimulation. J Physiol doi: 10.1113/jphysiol.2006.118711 (in press)
Saltiel P, Rossignol S (2004a) Critical points in the forelimb fictive locomotor cycle and motor coordination: evidence from the effects of tonic proprioceptive perturbations in the cat. J Neurophysiol 92:1329–1341
Saltiel P, Rossignol S (2004b) Critical points in the forelimb fictive locomotor cycle of the cat and motor coordination: effects of phasic retractions and protractions of the shoulder in the cat. J Neurophysiol 92:1342–1356
Schafer SS (1974) The discharge frequencies of primary muscle spindle endings during simultaneous stimulation of two fusimotor filaments. Pflugers Arch 350:359–372
Schmidt BJ, Meyers DER, Fleshman JL, Tokuriki M, Burke RE (1988) Phasic modulation of short latency cutaneous excitation in flexor digitorum longus motoneurons during fictive locomotion. Exp Brain Res 71:568–578
Schomburg ED, Petersen N, Barajon I, Hultborn H (1998) Flexor reflex afferents reset the step cycle during fictive locomotion in the cat. Exp Brain Res 122:339–350
Schouenborg J (2002) Modular organisation and spinal somatosensory imprinting. Brain Res Rev 40:80–91
Sharrard WJ (1964) The segmental innervation of the lower limb muscles in man. Annu R Coll Surg Engl 35:106–122
Sherrington CS (1910a) Flexion-reflex of the limb, crossed extension-reflex, and reflex stepping and standing. J Physiol 40:28–121
Sherrington CS (1910b) Remarks on the reflex mechanism of the step. Brain 33:1–25
Shik ML, Severin FV, Orlovsky GN (1966) Control of walking and running by means of electrical stimulation of the mid-brain. Biophysics 11:756–765
Sinkjaer T, Andersen JB, Ladouceur M, Christensen LOD, Nielsen J (2000) Major role for sensory feedback in soleus EMG activity in the stance phase of walking in man. J Physiol 523:817–827
Stein PS, Daniels-McQueen S (2002) Modular organization of turtle spinal interneurons during normal and deletion fictive rostral scratching. J Neurosci 22:6800–6809
Stein RB, Misiaszek JE, Pearson KG (2000) Functional role of muscle reflexes for force generation in the decerebrate walking cat. J Physiol 525:781–791
Stephens MJ, Yang JF (1999) Loading during the stance phase of walking in humans increases the extensor EMG amplitude but does not change the duration of the step cycle. Exp Brain Res 124:363–370
Taga G (1995a) A model of the neuro-musculo-skeletal system for human locomotion: I. Emergence of basic gait. Biol Cybern 73:97–111
Taga G (1995b) A model of the neuro-musculo-skeletal system for human locomotion. II Real-time adaptability under various constraints. Biol Cybern 73:113–121
Taga G (1998) A model of the neuro-musculo-skeletal system for anticipatory adjustment of human locomotion during obstacle avoidance. Biol Cybern 78:9–17
Taga G, Yamaguchi Y, Shimizu H (1991) Self-organized control of bipedal locomotion by neural oscillators in unpredictable environment. Biol Cybern 65:147–159
Taub E (1976) Motor behavior following deafferentation in the developing and motorically mature monkey. In: Herman R, Grillner S, Ralston HJ, Stein PSG, Stuart D (eds) Neural control of locomotion. Plenum, New York, pp 675-705
Taub E, Berman AJ (1968) Movement and learning in the absence of sensory feedback. In: Sandford, Frecdman J (eds) The neurophysiology of spatially oriented behaviour. Dorsey Press, Homewood, pp 173–191
Taylor A, Durbaba R, Ellaway PH (2004) Direct and indirect assessment of gamma-motor firing patterns. Can J Physiol Pharmacol 82:793–802
Twitchell TE (1954) Sensory factors in purposive movement. J Neurophysiol 17:239–252
Vanderhorst VGJM, Holstege G (1997) Organization of lumbosacral motoneuronal cell groups innervating hindlimb, pelvic floor, and axial muscles in the cat. J Comp Neurol 382: 46–76
Viala G, Buser P (1974) Inhibition des activités spinales à caractère locomoteur par une modalité particulière de stimulation somatique chez le lapin. Exp Brain Res 21:275–284
Viala G, Orsal D, Buser P (1978) Cutaneous fiber groups involved in the inhibition of fictive locomotion in the rabbit. Exp Brain Res 33:257–267
Wadden T, Ekeberg O (1998) A neuro-mechanical model of legged locomotion: single leg control. Biol Cybern 79: 161–173
Wallen P, Williams TL (1984) Fictive locomotion in lamprey spinal cord in vitro compared with swimming in the intact and spinal animal. J Physiol 347:225–239
Wand P, Prochazka A, Sontag KH (1980) Neuromuscular responses to gait perturbations in freely moving cats. Exp Brain Res 38:109–114
Wetzel MC, Atwater AE, Wait JV, Stuart DG (1976) Kinematics of locomotion by cats with a single hindlimb deafferented. J Neurophysiol 39:667–678
Wu JZ, Dong RG, Rakheja S, Schopper AW, Smutz WP (2004) A structural fingertip model for simulating of the biomechanics of tactile sensation. Med Eng Phys 26:165–175
Yakovenko S, Mushahwar V, VanderHorst V, Holstege G, Prochazka A (2002) Spatiotemporal activation of lumbosacral motoneurons in the locomotor step cycle. J Neurophysiol 87:1542–1553
Yakovenko S, Gritsenko V, Prochazka A (2004) Contribution of stretch reflexes to locomotor control: a modeling study. Biol Cybern 90:146–155
Yang JF, Stein RB, James KB (1991) Contribution of peripheral afferents to the activation of the soleus muscle during walking in humans. Exp Brain Res 87:679–687
Zajac FE (1989) Muscle and tendon: properties, models, scaling, and application to biomechanics and motor control. Crit Rev Biomed Eng 17:359–411
Zajac FE (2002) Understanding muscle coordination of the human leg with dynamical simulations. J Biomech 35:1011–1018
Zajac FE, Neptune RR, Kautz SA (2002) Biomechanics and muscle coordination of human walking: Part I. Introduction to concepts, power transfer, dynamics and simulations. Gait Posture 16:215–232
Zajac FE, Neptune RR, Kautz SA (2003) Biomechanics and muscle coordination of human walking: Part II. Lessons from dynamical simulations and clinical implications. Gait Posture 17:1–17
Zehr EP, Stein RB (1999) What functions do reflexes serve during human locomotion? Prog Neurobiol 58:185–205
Zehr EP, Komiyama T, Stein RB (1997) Cutaneous reflexes during human gait: electromyographic and kinematic responses to electrical stimulation. J Neurophysiol 77:3311–3325
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Frigon, A., Rossignol, S. Experiments and models of sensorimotor interactions during locomotion. Biol Cybern 95, 607–627 (2006). https://doi.org/10.1007/s00422-006-0129-x
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DOI: https://doi.org/10.1007/s00422-006-0129-x