Jean-pierre Gossard - Academia.edu (original) (raw)

Papers by Jean-pierre Gossard

Research paper thumbnail of Recovery of locomotion in cats after severe contusion of the low thoracic spinal cord

Journal of Neurophysiology

The recovery of quadrupedal treadmill locomotion after a large bilateral contusion at the low tho... more The recovery of quadrupedal treadmill locomotion after a large bilateral contusion at the low thoracic T10 spinal level and the ability to negotiate obstacles were studied for 5 wk in 16 cats. Ten cats were further completely spinalized at T13 and were found to walk with the hindlimbs within 24–72 h. We conclude that the extent of locomotor recovery after large spinal contusions hinges both on remnant supraspinal pathways and on a spinal pattern generator.

Research paper thumbnail of Ladder Treadmill: a method to assess locomotion in cats with an intact or lesioned spinal cord

The Journal of neuroscience : the official journal of the Society for Neuroscience, Jan 4, 2017

After lesions of the CNS, locomotor abilities of animals (mainly cats) are often assessed on a si... more After lesions of the CNS, locomotor abilities of animals (mainly cats) are often assessed on a simple flat treadmill (FTM) which imposes little demands on supraspinal structures as is the case when walking on targets. Therefore, the aims of the present work were 1) to develop a treadmill allowing the assessment of locomotion of intact cats required to place the paws on the rungs of a moving ladder treadmill (LTM); 2) to assess the capability of cats after a unilateral spinal hemisection at T10 to cope with such a demanding locomotor task; 3) to regularly train cats for 6wks on the LTM to see whether such regular training improves locomotor recovery on the FTM. A significant improvement would indicate that LTM training maximizes the contribution of spinal locomotor circuits as well as remnant supraspinal inputs. We used altogether 9 cats, 7 females and 2 males. Six were used to compare the EMG and kinematic locomotor characteristics during walking on the FTM and LTM. We found that th...

Research paper thumbnail of Plastic Changes in Lumbar Locomotor Networks after a Partial Spinal Cord Injury in Cats

The Journal of neuroscience : the official journal of the Society for Neuroscience, Jan 24, 2015

After an incomplete spinal cord injury (SCI), we know that plastic reorganization occurs in supra... more After an incomplete spinal cord injury (SCI), we know that plastic reorganization occurs in supraspinal structures with residual descending tracts. However, our knowledge about spinal plasticity is rather limited. Our recent studies point to changes within the spinal cord below the lesion. After a lateral left hemisection (T10), cats recovered stepping with both hindlimbs within 3 weeks. After a complete section (T13) in these cats, bilateral stepping was seen on the next day, a skill usually acquired after several weeks of treadmill training. This indicates that durable plastic changes occurred below the lesion. However, because sensory feedback entrains the stepping rhythm, it is difficult to reveal central pattern generator (CPG) adaptation. Here, we investigated whether lumbar segments of cats with a chronic hemisection were able to generate fictive locomotion-that is, without phasic sensory feedback as monitored by five muscle nerves in each hindlimb. With a chronic left hemise...

Research paper thumbnail of The "beneficial" effects of locomotor training after various types of spinal lesions in cats and rats

Progress in brain research, 2015

This chapter reviews a number of experiments on the recovery of locomotion after various types of... more This chapter reviews a number of experiments on the recovery of locomotion after various types of spinal lesions and locomotor training mainly in cats. We first recall the major evidence on the recovery of hindlimb locomotion in completely spinalized cats at the T13 level and the role played by the spinal locomotor network, also known as the central pattern generator, as well as the beneficial effects of locomotor training on this recovery. Having established that hindlimb locomotion can recover, we raise the issue as to whether spinal plastic changes could also contribute to the recovery after partial spinal lesions such as unilateral hemisections. We found that after such hemisection at T10, cats could recover quadrupedal locomotion and that deficits could be improved by training. We further showed that, after a complete spinalization a few segments below the first hemisection (at T13, i.e., the level of previous studies on spinalization), cats could readily walk with the hindlimb...

Research paper thumbnail of Task-dependent presynaptic inhibition

The Journal of neuroscience : the official journal of the Society for Neuroscience, 2003

This study compares the level of presynaptic inhibition during two rhythmic movements in the cat:... more This study compares the level of presynaptic inhibition during two rhythmic movements in the cat: locomotion and scratch. Dorsal rootlets from L6, L7, or S1 segments were cut, and their proximal stumps were recorded during fictive locomotion occurring spontaneously in decerebrate cats and during fictive scratch induced by d-tubocurarine applied on the C1 and C2 segments. Compared with rest, the number of antidromic spikes was increased (by 12%) during locomotion, whereas it was greatly decreased (31%) during scratch, and the amplitude of dorsal root potentials (DRPs), evoked by stimulating a muscle nerve, was slightly decreased (7%) during locomotion but much more so during scratch (53%). When compared with locomotion, the decrease in the number of antidromic spikes (45%) and the decrease in DRP amplitude (43%) during scratch were of similar magnitude. Also, the amplitude of primary afferent depolarization (PAD), recorded with micropipettes in axons (n = 13) of two cats, was found t...

Research paper thumbnail of Spinal cats on the treadmill: changes in load pathways

The Journal of neuroscience : the official journal of the Society for Neuroscience, 2003

Treadmill training and clonidine, an alpha-2 noradrenergic agonist, have been shown to improve lo... more Treadmill training and clonidine, an alpha-2 noradrenergic agonist, have been shown to improve locomotion after spinal cord injury. We speculate that transmission in load pathways, which are involved in body support during stance, is specifically modified by training. This was evaluated by comparing two groups of spinal cats; one group (n = 11) was trained to walk until full-weight-bearing (3-4 weeks), and the other (shams; n = 7) was not. During an acute experiment, changes in group I pathways, monosynaptic excitation, disynaptic inhibition, and polysynaptic excitation were investigated by measuring the response amplitude in extensor motoneurons before and after clonidine injection. Monosynaptic excitation was not modified by clonidine but was decreased significantly by training. Disynaptic inhibition was significantly decreased by clonidine in both groups, but more significantly in trained cats, and significantly reduced by training after clonidine. Also, clonidine could reverse g...

Research paper thumbnail of Preface

Progress in Brain Research, 2010

Research paper thumbnail of Preface. Breathe, walk and chew: the neural challenge: part II

Progress in brain research, 2011

Research paper thumbnail of In memoriam: James P. Lund

Progress in brain research, 2011

Research paper thumbnail of Key mechanisms for setting the input–output gain across the motoneuron pool

Progress in Brain Research, 2004

This chapter summarizes a number of factors that control the &amp... more This chapter summarizes a number of factors that control the "input-output" function across the motoneurons (MNs) comprising a single spinal motor nucleus. The main focus is on intrinsic properties of individual MNs that can be controlled by neuromodulators. These include: (1) amplification of the synaptic input at the cell's dendritic level by voltage-gated, persistent inward currents (plateau potentials); and (2) transduction of the net synaptic excitation into a frequency code (the MN's stimulus current-spike frequency relation) at the cell's soma/initial segment. Two other aspects of the synaptic control of MNs, which may affect their input-output gain, are also discussed. They include the hypotheses that: (1) a non-uniform distribution of synaptic effects to low- and high-threshold motor units causes a change in recruitment gain; and (2) recurrent inhibition, via motor axon collaterals and Renshaw cells, functions as a variable gain regulator of MN discharge.

Research paper thumbnail of Bulbospinal control of spinal cord pathways generating locomotor extensor activities in the cat

The Journal of Physiology, 2000

Research paper thumbnail of Asymmetric control of cycle period by the spinal locomotor rhythm generator in the adult cat

The Journal of Physiology, 2009

During walking, a change in speed is accomplished by varying the duration of the stance phase, wh... more During walking, a change in speed is accomplished by varying the duration of the stance phase, while the swing phase remains relatively invariant. To determine if this asymmetry in the control of locomotor cycles is an inherent property of the spinal central pattern generator (CPG), we recorded episodes of fictive locomotion in decerebrate cats with or without a complete spinal transection (acute or chronic). During fictive locomotion, stance and swing phases typically correspond to extension and flexion phases, respectively. The extension and flexion phases were determined by measuring the duration of extensor and flexor bursts, respectively. In the vast majority of locomotor episodes, cycle period varied more with the extension phase. This was found without phasic sensory feedback, supraspinal structures, pharmacology or sustained stimulation. We conclude that the control of walking speed is governed by an asymmetry within the organization of the spinal CPG, which can be modified by extraneous factors.

Research paper thumbnail of Independent Control of Presynaptic Inhibition by Reticulospinal and Sensory Inputs at Rest and during Rhythmic Activities in the Cat

Journal of Neuroscience, 2013

To be functionally relevant during movement, the transmission from primary afferents must be effi... more To be functionally relevant during movement, the transmission from primary afferents must be efficiently controlled by presynaptic inhibition. Sensory feedback, central pattern generators, and supraspinal structures can all evoke presynaptic inhibition, but we do not understand how these inputs interact during movement. Here, we investigated the convergence of inputs from the reticular formation and sensory afferents on presynaptic inhibitory pathways and their modulation at rest and during two fictive motor tasks (locomotion and scratch) in decerebrate cats. The amplitude of primary afferent depolarization (PAD), an estimate of presynaptic inhibition, was recorded in individual afferents with intra-axonal recordings and in a mix of afferents in lumbar dorsal rootlets (dorsal root potential [DRP]) with bipolar electrodes. There was no spatial facilitation between inputs from reticulospinal and sensory afferents with DRPs or PADs, indicating an absence of convergence. However, spatial facilitation could be observed by combining two sensory inputs, indicating that convergence was possible. Task-dependent changes in the amplitude of responses were similar for reticulospinal and sensory inputs, increasing during fictive locomotion and decreasing during fictive scratch. During fictive locomotion, DRP and PAD amplitudes evoked by reticulospinal inputs were increased during the flexion phase, whereas sensory-evoked DRPs and PADs showed maximal amplitude in either flexion or extension phases. During fictive scratch, the amplitudes of DRPs and PADs evoked by both sources were maximal in flexion. The absence of spatial facilitation and different phase-dependent modulation patterns during fictive locomotion are consistent with independent presynaptic inhibitory pathways for reticulospinal and sensory inputs.

Research paper thumbnail of Evidence for Specialized Rhythm-Generating Mechanisms in the Adult Mammalian Spinal Cord

Journal of Neuroscience, 2010

Locomotion and scratch are characterized by alternation of flexion and extension phases within on... more Locomotion and scratch are characterized by alternation of flexion and extension phases within one hindlimb, which are mediated by rhythm-generating circuitry within the spinal cord. By definition, the rhythm generator controls cycle period, phase durations, and phase transitions. The aim was to determine whether rhythm-generating mechanisms for locomotion and scratch are similar in adult decerebrate cats. The regulation of cycle period during fictive scratching was evaluated, as were the effects of specific sensory inputs on phase durations and transitions during spontaneous fictive locomotion and pinna-evoked fictive scratching. Results show that cycle period during fictive scratching varied predominantly with flexion phase duration, contrary to spontaneous fictive locomotion, where cycle period varied with extension phase duration. Ankle dorsiflexion greatly increased extension phase duration and cycle period during fictive locomotion but did not alter cycle period during scratching. Moreover, stimulating the plantaris (ankle extensor muscle) nerve during flexion reset the locomotor rhythm to extension but not the scratch rhythm. Stimulating the plantaris nerve during extension prolonged the extension phase and cycle period during fictive locomotion but not during fictive scratching. Stimulating the sartorius nerve (hip flexor muscle) during early flexion reduced the flexion phase and cycle period during fictive locomotion, but considerably prolonged the flexion phase and cycle period during fictive scratching. These data indicate that cycle period, phase durations, and phase transitions are not regulated similarly during fictive locomotion and scratching, with or without sensory inputs, providing evidence for specialized rhythm-generating mechanisms within the adult mammalian spinal cord.

Research paper thumbnail of Dynamic Sensorimotor Interactions in Locomotion

Physiological Reviews, 2006

Locomotion results from intricate dynamic interactions between a central program and feedback mec... more Locomotion results from intricate dynamic interactions between a central program and feedback mechanisms. The central program relies fundamentally on a genetically determined spinal circuitry (central pattern generator) capable of generating the basic locomotor pattern and on various descending pathways that can trigger, stop, and steer locomotion. The feedback originates from muscles and skin afferents as well as from special senses (vision, audition, vestibular) and dynamically adapts the locomotor pattern to the requirements of the environment. The dynamic interactions are ensured by modulating transmission in locomotor pathways in a state- and phase-dependent manner. For instance, proprioceptive inputs from extensors can, during stance, adjust the timing and amplitude of muscle activities of the limbs to the speed of locomotion but be silenced during the opposite phase of the cycle. Similarly, skin afferents participate predominantly in the correction of limb and foot placement ...

Research paper thumbnail of Effects of Ankle and Hip Muscle Afferent Inputs on Rhythm Generation During Fictive Locomotion

Journal of Neurophysiology, 2010

Hip position and loading of limb extensors are major sensory cues for the initiation and duration... more Hip position and loading of limb extensors are major sensory cues for the initiation and duration of different phases during walking. Although these inputs have pathways projecting to the locomotor rhythm generator, their effects may vary in different parts of the locomotor cycle. In the present study, the plantaris (Pl), sartorius (Sart), rectus femoris (RF), and caudal gluteal (cGlu) nerves were stimulated at group I and/or group II strength during spontaneous fictive locomotion in 16 adult decerebrate cats. These nerves supply muscles that extend the ankle (Pl), flex the hip (Sart, RF), or extend the hip (cGlu). Stimuli were given at six epochs of the locomotor cycle to evaluate when they access the rhythm generator. Group I afferents from Pl nerve always reset the locomotor rhythm; stimulation during extension prolonged cycle period and extension phase duration, while stimulation during flexion terminated flexion and initiated extension. On the other hand, stimulating RF and cGl...

Research paper thumbnail of Corticospinal control of locomotor pathways generating extensor activities in the cat

Experimental Brain Research, 2001

Interneuronal convergence of corticospinal and segmental pathways involved with the generation of... more Interneuronal convergence of corticospinal and segmental pathways involved with the generation of extensor activities during locomotion was investigated in decerebrate and partially spinalized cats. L-dihydroxyphenylalanine (L-DOPA) was slowly injected until long-latency, long-lasting discharges could be evoked by the stimulation of contralateral flexor reflex afferents (coFRA) and the group I autogenetic inhibition was reversed to polysynaptic excitation in extensor motoneurons. Under these conditions, we stimulated in alternation the contralateral pyramidal tract (PT), group I afferents from knee and ankle extensor muscles, and both stimuli together. We did the same for the stimulation of PT and of coFRA. Clear polysynaptic EPSPs could be evoked from all three sources in 32 extensor motoneurons. Convergence was inferred from spatial facilitation, which occurred when the amplitude of the EPSPs evoked by the combined stimuli was notably larger than the algebraic sum of the EPSPs evoked by individual stimulation. Spatial facilitation was found between PT and extensor group I inputs in 30/59 tests (51%) in 20 motoneurons and in all cases (6/6) between PT and coFRA in six motoneurons. When fictive locomotion was induced with further injection of L-DOPA, PT descending volleys from the same stimulating site could reset the stepping rhythm by initiating bursts of activity in all extensors. These results indicate that at least some of the corticospinal fibers project onto interneurons shared by the coFRA and the polysynaptic excitatory group I pathways to extensors. The implications of such convergence patterns on the organization of the extensor "halfcenter" for locomotion are discussed.

Research paper thumbnail of Modulation of monosynaptic transmission by presynaptic inhibition during fictive locomotion in the cat

Brain Research, 2003

The effect of multisensory inputs onto the presynaptic inhibitory pathways affecting IA terminals... more The effect of multisensory inputs onto the presynaptic inhibitory pathways affecting IA terminals was studied during fictive locomotion in decerebrated cats. The effect was evaluated from changes in amplitude of the monosynaptic excitatory postsynaptic potential (EPSP) measured in lumbosacral motoneurones. Responses were grouped and averaged according to their timing within the step cycle divided into five bins. Presynaptic inhibition was evoked by stimulating group I afferents from the posterior biceps-semitendinosus (PBSt) muscles and one of three cutaneous nerves: superficial peroneal (SP), sural and saphenous. Statistical analysis was applied to compare (1) EPSPs conditioned by PBSt input alone and those conditioned by the combined PBSt and cutaneous inputs, and (2) each bin dividing the step cycle to disclose phase-dependent changes. Results from 19 motoneurones showed that: (1) there was a significant phase-dependent modulation in EPSP amplitude (by 25%) with the maximum usually occurring during the depolarized phase; (2) PBSt alone reduced the EPSP amplitude (by 21%) in 3.2 bins on average; (3) combined PBSt and cutaneous stimuli further modified (up or down) the EPSP amplitude in half the trials but only in one to two bins; and (4) the most efficient cutaneous nerve (SP) usually decreased the PBSt-evoked reduction in EPSP size. Minimal changes in membrane input resistance suggest that the EPSP modifications were mostly due to presynaptic inhibition. Results indicate that muscle afferents can induce an important phase-dependent presynaptic inhibition of monosynaptic transmission and that concomitant activation of cutaneous afferents can alter this inhibition but only for a restricted part of the step cycle.

Research paper thumbnail of The effects of antidromic discharges on orthodromic firing of primary afferents in the cat

Brain Research, 1999

This study investigated the effects of antidromically conducted nerve impulses on the transmissio... more This study investigated the effects of antidromically conducted nerve impulses on the transmission of orthodromic volleys in primary Ž. Ž. afferents of the hindlimb in decerebrated paralyzed cats. Two protocols were used: A Single skin and muscle afferents N s 20 isolated Ž. from the distal part of cut dorsal rootlets L7-S1 were recorded while stimulation was applied more caudally. The results showed that during the trains of three to 20 stimuli, the orthodromic firing frequency decreased or ceased, depending on the frequency of stimulation. Ž. Remarkably, subsequent to these trains, the occurrence of orthodromic spikes could be delayed for hundreds of ms 15r20 afferents and Ž. sometimes stopped for several seconds 10r20 afferents. Longer stimulation trains, simulating antidromic bursts reported during Ž. locomotion, caused a progressive decrease, and a slow recovery of, orthodromic firing frequency 7r20 afferents , indicating a Ž. cumulative long-lasting depressing effect from successive bursts. B Identified stretch-sensitive muscle afferents were recorded intra-axonally and antidromic spikes were evoked by the injection of square pulses of current through the micropipette. In this case, one to three antidromic spikes were sufficient to delay the occurrence of the next orthodromic spike by more than one control inter-spike interval. If the control inter-spike interval was decreased by stretching the muscle, the delay evoked by antidromic spikes decreased proportionally. Overall, these findings suggest that antidromic activity could alter the mechanisms underlying spike generation in peripheral sensory receptors and modify the orthodromic discharges of afferents during locomotion.

Research paper thumbnail of Phase-dependent modulation of primary afferent depolarization in single cutaneous primary afferents evoked by peripheral stimulation during fictive locomotion in the cat

Brain Research, 1990

... The results thus show that the transmission in PAD pathways activatedby cutaneous inputs ... ... more ... The results thus show that the transmission in PAD pathways activatedby cutaneous inputs ... standarddeviations as for SP units.20F--Z0 00 0MIN *MAX o000000OOOOOO o ... is unknown, we may speculate for discussionpurposes (see Introduction), that rhythmic changes ofafferent ...

Research paper thumbnail of Recovery of locomotion in cats after severe contusion of the low thoracic spinal cord

Journal of Neurophysiology

The recovery of quadrupedal treadmill locomotion after a large bilateral contusion at the low tho... more The recovery of quadrupedal treadmill locomotion after a large bilateral contusion at the low thoracic T10 spinal level and the ability to negotiate obstacles were studied for 5 wk in 16 cats. Ten cats were further completely spinalized at T13 and were found to walk with the hindlimbs within 24–72 h. We conclude that the extent of locomotor recovery after large spinal contusions hinges both on remnant supraspinal pathways and on a spinal pattern generator.

Research paper thumbnail of Ladder Treadmill: a method to assess locomotion in cats with an intact or lesioned spinal cord

The Journal of neuroscience : the official journal of the Society for Neuroscience, Jan 4, 2017

After lesions of the CNS, locomotor abilities of animals (mainly cats) are often assessed on a si... more After lesions of the CNS, locomotor abilities of animals (mainly cats) are often assessed on a simple flat treadmill (FTM) which imposes little demands on supraspinal structures as is the case when walking on targets. Therefore, the aims of the present work were 1) to develop a treadmill allowing the assessment of locomotion of intact cats required to place the paws on the rungs of a moving ladder treadmill (LTM); 2) to assess the capability of cats after a unilateral spinal hemisection at T10 to cope with such a demanding locomotor task; 3) to regularly train cats for 6wks on the LTM to see whether such regular training improves locomotor recovery on the FTM. A significant improvement would indicate that LTM training maximizes the contribution of spinal locomotor circuits as well as remnant supraspinal inputs. We used altogether 9 cats, 7 females and 2 males. Six were used to compare the EMG and kinematic locomotor characteristics during walking on the FTM and LTM. We found that th...

Research paper thumbnail of Plastic Changes in Lumbar Locomotor Networks after a Partial Spinal Cord Injury in Cats

The Journal of neuroscience : the official journal of the Society for Neuroscience, Jan 24, 2015

After an incomplete spinal cord injury (SCI), we know that plastic reorganization occurs in supra... more After an incomplete spinal cord injury (SCI), we know that plastic reorganization occurs in supraspinal structures with residual descending tracts. However, our knowledge about spinal plasticity is rather limited. Our recent studies point to changes within the spinal cord below the lesion. After a lateral left hemisection (T10), cats recovered stepping with both hindlimbs within 3 weeks. After a complete section (T13) in these cats, bilateral stepping was seen on the next day, a skill usually acquired after several weeks of treadmill training. This indicates that durable plastic changes occurred below the lesion. However, because sensory feedback entrains the stepping rhythm, it is difficult to reveal central pattern generator (CPG) adaptation. Here, we investigated whether lumbar segments of cats with a chronic hemisection were able to generate fictive locomotion-that is, without phasic sensory feedback as monitored by five muscle nerves in each hindlimb. With a chronic left hemise...

Research paper thumbnail of The "beneficial" effects of locomotor training after various types of spinal lesions in cats and rats

Progress in brain research, 2015

This chapter reviews a number of experiments on the recovery of locomotion after various types of... more This chapter reviews a number of experiments on the recovery of locomotion after various types of spinal lesions and locomotor training mainly in cats. We first recall the major evidence on the recovery of hindlimb locomotion in completely spinalized cats at the T13 level and the role played by the spinal locomotor network, also known as the central pattern generator, as well as the beneficial effects of locomotor training on this recovery. Having established that hindlimb locomotion can recover, we raise the issue as to whether spinal plastic changes could also contribute to the recovery after partial spinal lesions such as unilateral hemisections. We found that after such hemisection at T10, cats could recover quadrupedal locomotion and that deficits could be improved by training. We further showed that, after a complete spinalization a few segments below the first hemisection (at T13, i.e., the level of previous studies on spinalization), cats could readily walk with the hindlimb...

Research paper thumbnail of Task-dependent presynaptic inhibition

The Journal of neuroscience : the official journal of the Society for Neuroscience, 2003

This study compares the level of presynaptic inhibition during two rhythmic movements in the cat:... more This study compares the level of presynaptic inhibition during two rhythmic movements in the cat: locomotion and scratch. Dorsal rootlets from L6, L7, or S1 segments were cut, and their proximal stumps were recorded during fictive locomotion occurring spontaneously in decerebrate cats and during fictive scratch induced by d-tubocurarine applied on the C1 and C2 segments. Compared with rest, the number of antidromic spikes was increased (by 12%) during locomotion, whereas it was greatly decreased (31%) during scratch, and the amplitude of dorsal root potentials (DRPs), evoked by stimulating a muscle nerve, was slightly decreased (7%) during locomotion but much more so during scratch (53%). When compared with locomotion, the decrease in the number of antidromic spikes (45%) and the decrease in DRP amplitude (43%) during scratch were of similar magnitude. Also, the amplitude of primary afferent depolarization (PAD), recorded with micropipettes in axons (n = 13) of two cats, was found t...

Research paper thumbnail of Spinal cats on the treadmill: changes in load pathways

The Journal of neuroscience : the official journal of the Society for Neuroscience, 2003

Treadmill training and clonidine, an alpha-2 noradrenergic agonist, have been shown to improve lo... more Treadmill training and clonidine, an alpha-2 noradrenergic agonist, have been shown to improve locomotion after spinal cord injury. We speculate that transmission in load pathways, which are involved in body support during stance, is specifically modified by training. This was evaluated by comparing two groups of spinal cats; one group (n = 11) was trained to walk until full-weight-bearing (3-4 weeks), and the other (shams; n = 7) was not. During an acute experiment, changes in group I pathways, monosynaptic excitation, disynaptic inhibition, and polysynaptic excitation were investigated by measuring the response amplitude in extensor motoneurons before and after clonidine injection. Monosynaptic excitation was not modified by clonidine but was decreased significantly by training. Disynaptic inhibition was significantly decreased by clonidine in both groups, but more significantly in trained cats, and significantly reduced by training after clonidine. Also, clonidine could reverse g...

Research paper thumbnail of Preface

Progress in Brain Research, 2010

Research paper thumbnail of Preface. Breathe, walk and chew: the neural challenge: part II

Progress in brain research, 2011

Research paper thumbnail of In memoriam: James P. Lund

Progress in brain research, 2011

Research paper thumbnail of Key mechanisms for setting the input–output gain across the motoneuron pool

Progress in Brain Research, 2004

This chapter summarizes a number of factors that control the &amp... more This chapter summarizes a number of factors that control the "input-output" function across the motoneurons (MNs) comprising a single spinal motor nucleus. The main focus is on intrinsic properties of individual MNs that can be controlled by neuromodulators. These include: (1) amplification of the synaptic input at the cell's dendritic level by voltage-gated, persistent inward currents (plateau potentials); and (2) transduction of the net synaptic excitation into a frequency code (the MN's stimulus current-spike frequency relation) at the cell's soma/initial segment. Two other aspects of the synaptic control of MNs, which may affect their input-output gain, are also discussed. They include the hypotheses that: (1) a non-uniform distribution of synaptic effects to low- and high-threshold motor units causes a change in recruitment gain; and (2) recurrent inhibition, via motor axon collaterals and Renshaw cells, functions as a variable gain regulator of MN discharge.

Research paper thumbnail of Bulbospinal control of spinal cord pathways generating locomotor extensor activities in the cat

The Journal of Physiology, 2000

Research paper thumbnail of Asymmetric control of cycle period by the spinal locomotor rhythm generator in the adult cat

The Journal of Physiology, 2009

During walking, a change in speed is accomplished by varying the duration of the stance phase, wh... more During walking, a change in speed is accomplished by varying the duration of the stance phase, while the swing phase remains relatively invariant. To determine if this asymmetry in the control of locomotor cycles is an inherent property of the spinal central pattern generator (CPG), we recorded episodes of fictive locomotion in decerebrate cats with or without a complete spinal transection (acute or chronic). During fictive locomotion, stance and swing phases typically correspond to extension and flexion phases, respectively. The extension and flexion phases were determined by measuring the duration of extensor and flexor bursts, respectively. In the vast majority of locomotor episodes, cycle period varied more with the extension phase. This was found without phasic sensory feedback, supraspinal structures, pharmacology or sustained stimulation. We conclude that the control of walking speed is governed by an asymmetry within the organization of the spinal CPG, which can be modified by extraneous factors.

Research paper thumbnail of Independent Control of Presynaptic Inhibition by Reticulospinal and Sensory Inputs at Rest and during Rhythmic Activities in the Cat

Journal of Neuroscience, 2013

To be functionally relevant during movement, the transmission from primary afferents must be effi... more To be functionally relevant during movement, the transmission from primary afferents must be efficiently controlled by presynaptic inhibition. Sensory feedback, central pattern generators, and supraspinal structures can all evoke presynaptic inhibition, but we do not understand how these inputs interact during movement. Here, we investigated the convergence of inputs from the reticular formation and sensory afferents on presynaptic inhibitory pathways and their modulation at rest and during two fictive motor tasks (locomotion and scratch) in decerebrate cats. The amplitude of primary afferent depolarization (PAD), an estimate of presynaptic inhibition, was recorded in individual afferents with intra-axonal recordings and in a mix of afferents in lumbar dorsal rootlets (dorsal root potential [DRP]) with bipolar electrodes. There was no spatial facilitation between inputs from reticulospinal and sensory afferents with DRPs or PADs, indicating an absence of convergence. However, spatial facilitation could be observed by combining two sensory inputs, indicating that convergence was possible. Task-dependent changes in the amplitude of responses were similar for reticulospinal and sensory inputs, increasing during fictive locomotion and decreasing during fictive scratch. During fictive locomotion, DRP and PAD amplitudes evoked by reticulospinal inputs were increased during the flexion phase, whereas sensory-evoked DRPs and PADs showed maximal amplitude in either flexion or extension phases. During fictive scratch, the amplitudes of DRPs and PADs evoked by both sources were maximal in flexion. The absence of spatial facilitation and different phase-dependent modulation patterns during fictive locomotion are consistent with independent presynaptic inhibitory pathways for reticulospinal and sensory inputs.

Research paper thumbnail of Evidence for Specialized Rhythm-Generating Mechanisms in the Adult Mammalian Spinal Cord

Journal of Neuroscience, 2010

Locomotion and scratch are characterized by alternation of flexion and extension phases within on... more Locomotion and scratch are characterized by alternation of flexion and extension phases within one hindlimb, which are mediated by rhythm-generating circuitry within the spinal cord. By definition, the rhythm generator controls cycle period, phase durations, and phase transitions. The aim was to determine whether rhythm-generating mechanisms for locomotion and scratch are similar in adult decerebrate cats. The regulation of cycle period during fictive scratching was evaluated, as were the effects of specific sensory inputs on phase durations and transitions during spontaneous fictive locomotion and pinna-evoked fictive scratching. Results show that cycle period during fictive scratching varied predominantly with flexion phase duration, contrary to spontaneous fictive locomotion, where cycle period varied with extension phase duration. Ankle dorsiflexion greatly increased extension phase duration and cycle period during fictive locomotion but did not alter cycle period during scratching. Moreover, stimulating the plantaris (ankle extensor muscle) nerve during flexion reset the locomotor rhythm to extension but not the scratch rhythm. Stimulating the plantaris nerve during extension prolonged the extension phase and cycle period during fictive locomotion but not during fictive scratching. Stimulating the sartorius nerve (hip flexor muscle) during early flexion reduced the flexion phase and cycle period during fictive locomotion, but considerably prolonged the flexion phase and cycle period during fictive scratching. These data indicate that cycle period, phase durations, and phase transitions are not regulated similarly during fictive locomotion and scratching, with or without sensory inputs, providing evidence for specialized rhythm-generating mechanisms within the adult mammalian spinal cord.

Research paper thumbnail of Dynamic Sensorimotor Interactions in Locomotion

Physiological Reviews, 2006

Locomotion results from intricate dynamic interactions between a central program and feedback mec... more Locomotion results from intricate dynamic interactions between a central program and feedback mechanisms. The central program relies fundamentally on a genetically determined spinal circuitry (central pattern generator) capable of generating the basic locomotor pattern and on various descending pathways that can trigger, stop, and steer locomotion. The feedback originates from muscles and skin afferents as well as from special senses (vision, audition, vestibular) and dynamically adapts the locomotor pattern to the requirements of the environment. The dynamic interactions are ensured by modulating transmission in locomotor pathways in a state- and phase-dependent manner. For instance, proprioceptive inputs from extensors can, during stance, adjust the timing and amplitude of muscle activities of the limbs to the speed of locomotion but be silenced during the opposite phase of the cycle. Similarly, skin afferents participate predominantly in the correction of limb and foot placement ...

Research paper thumbnail of Effects of Ankle and Hip Muscle Afferent Inputs on Rhythm Generation During Fictive Locomotion

Journal of Neurophysiology, 2010

Hip position and loading of limb extensors are major sensory cues for the initiation and duration... more Hip position and loading of limb extensors are major sensory cues for the initiation and duration of different phases during walking. Although these inputs have pathways projecting to the locomotor rhythm generator, their effects may vary in different parts of the locomotor cycle. In the present study, the plantaris (Pl), sartorius (Sart), rectus femoris (RF), and caudal gluteal (cGlu) nerves were stimulated at group I and/or group II strength during spontaneous fictive locomotion in 16 adult decerebrate cats. These nerves supply muscles that extend the ankle (Pl), flex the hip (Sart, RF), or extend the hip (cGlu). Stimuli were given at six epochs of the locomotor cycle to evaluate when they access the rhythm generator. Group I afferents from Pl nerve always reset the locomotor rhythm; stimulation during extension prolonged cycle period and extension phase duration, while stimulation during flexion terminated flexion and initiated extension. On the other hand, stimulating RF and cGl...

Research paper thumbnail of Corticospinal control of locomotor pathways generating extensor activities in the cat

Experimental Brain Research, 2001

Interneuronal convergence of corticospinal and segmental pathways involved with the generation of... more Interneuronal convergence of corticospinal and segmental pathways involved with the generation of extensor activities during locomotion was investigated in decerebrate and partially spinalized cats. L-dihydroxyphenylalanine (L-DOPA) was slowly injected until long-latency, long-lasting discharges could be evoked by the stimulation of contralateral flexor reflex afferents (coFRA) and the group I autogenetic inhibition was reversed to polysynaptic excitation in extensor motoneurons. Under these conditions, we stimulated in alternation the contralateral pyramidal tract (PT), group I afferents from knee and ankle extensor muscles, and both stimuli together. We did the same for the stimulation of PT and of coFRA. Clear polysynaptic EPSPs could be evoked from all three sources in 32 extensor motoneurons. Convergence was inferred from spatial facilitation, which occurred when the amplitude of the EPSPs evoked by the combined stimuli was notably larger than the algebraic sum of the EPSPs evoked by individual stimulation. Spatial facilitation was found between PT and extensor group I inputs in 30/59 tests (51%) in 20 motoneurons and in all cases (6/6) between PT and coFRA in six motoneurons. When fictive locomotion was induced with further injection of L-DOPA, PT descending volleys from the same stimulating site could reset the stepping rhythm by initiating bursts of activity in all extensors. These results indicate that at least some of the corticospinal fibers project onto interneurons shared by the coFRA and the polysynaptic excitatory group I pathways to extensors. The implications of such convergence patterns on the organization of the extensor "halfcenter" for locomotion are discussed.

Research paper thumbnail of Modulation of monosynaptic transmission by presynaptic inhibition during fictive locomotion in the cat

Brain Research, 2003

The effect of multisensory inputs onto the presynaptic inhibitory pathways affecting IA terminals... more The effect of multisensory inputs onto the presynaptic inhibitory pathways affecting IA terminals was studied during fictive locomotion in decerebrated cats. The effect was evaluated from changes in amplitude of the monosynaptic excitatory postsynaptic potential (EPSP) measured in lumbosacral motoneurones. Responses were grouped and averaged according to their timing within the step cycle divided into five bins. Presynaptic inhibition was evoked by stimulating group I afferents from the posterior biceps-semitendinosus (PBSt) muscles and one of three cutaneous nerves: superficial peroneal (SP), sural and saphenous. Statistical analysis was applied to compare (1) EPSPs conditioned by PBSt input alone and those conditioned by the combined PBSt and cutaneous inputs, and (2) each bin dividing the step cycle to disclose phase-dependent changes. Results from 19 motoneurones showed that: (1) there was a significant phase-dependent modulation in EPSP amplitude (by 25%) with the maximum usually occurring during the depolarized phase; (2) PBSt alone reduced the EPSP amplitude (by 21%) in 3.2 bins on average; (3) combined PBSt and cutaneous stimuli further modified (up or down) the EPSP amplitude in half the trials but only in one to two bins; and (4) the most efficient cutaneous nerve (SP) usually decreased the PBSt-evoked reduction in EPSP size. Minimal changes in membrane input resistance suggest that the EPSP modifications were mostly due to presynaptic inhibition. Results indicate that muscle afferents can induce an important phase-dependent presynaptic inhibition of monosynaptic transmission and that concomitant activation of cutaneous afferents can alter this inhibition but only for a restricted part of the step cycle.

Research paper thumbnail of The effects of antidromic discharges on orthodromic firing of primary afferents in the cat

Brain Research, 1999

This study investigated the effects of antidromically conducted nerve impulses on the transmissio... more This study investigated the effects of antidromically conducted nerve impulses on the transmission of orthodromic volleys in primary Ž. Ž. afferents of the hindlimb in decerebrated paralyzed cats. Two protocols were used: A Single skin and muscle afferents N s 20 isolated Ž. from the distal part of cut dorsal rootlets L7-S1 were recorded while stimulation was applied more caudally. The results showed that during the trains of three to 20 stimuli, the orthodromic firing frequency decreased or ceased, depending on the frequency of stimulation. Ž. Remarkably, subsequent to these trains, the occurrence of orthodromic spikes could be delayed for hundreds of ms 15r20 afferents and Ž. sometimes stopped for several seconds 10r20 afferents. Longer stimulation trains, simulating antidromic bursts reported during Ž. locomotion, caused a progressive decrease, and a slow recovery of, orthodromic firing frequency 7r20 afferents , indicating a Ž. cumulative long-lasting depressing effect from successive bursts. B Identified stretch-sensitive muscle afferents were recorded intra-axonally and antidromic spikes were evoked by the injection of square pulses of current through the micropipette. In this case, one to three antidromic spikes were sufficient to delay the occurrence of the next orthodromic spike by more than one control inter-spike interval. If the control inter-spike interval was decreased by stretching the muscle, the delay evoked by antidromic spikes decreased proportionally. Overall, these findings suggest that antidromic activity could alter the mechanisms underlying spike generation in peripheral sensory receptors and modify the orthodromic discharges of afferents during locomotion.

Research paper thumbnail of Phase-dependent modulation of primary afferent depolarization in single cutaneous primary afferents evoked by peripheral stimulation during fictive locomotion in the cat

Brain Research, 1990

... The results thus show that the transmission in PAD pathways activatedby cutaneous inputs ... ... more ... The results thus show that the transmission in PAD pathways activatedby cutaneous inputs ... standarddeviations as for SP units.20F--Z0 00 0MIN *MAX o000000OOOOOO o ... is unknown, we may speculate for discussionpurposes (see Introduction), that rhythmic changes ofafferent ...