The role of Renshaw cells in locomotion: antagonism of their excitation from motor axon collaterals with intravenous mecamylamine

Summary The contribution of Renshaw cell (RC) activity to the production of fictive locomotion in the mesencephalic preparation was examined using the nicotinic antagonist mecamylamine (MEC). After the i.v. administration of 3 doses of MEC (1.0 mg/kg) the following observations were made: 1) ventral root (VR) evoked discharge of RCs was decreased by up to 87.7%, 2) recurrent inhibitory postsynaptic potentials recorded in alpha motoneurons were greatly reduced or abolished, and 3) the rhythmic firing of RCs during the fictive step cycle was abolished in 83% of the cells examined. Locomotor drive potentials (LDPs) in motoneurons persisted during the fictive step cycle after MEC administration. Bursts of motoneuron firing during each fictive step cycle were characterized by increased frequency and number of spikes after MEC, although the burst duration was unaltered for similar step cycle lengths. A greater number and frequency of spikes per burst was also observed in Ia inhibitory interneurons (IaINs), which remained rhythmically active after MEC administration. It is concluded that Renshaw cells are not an integral part of the spinal central pattern generator for locomotion, nor do they control the timing of the motoneuron or IaIN bursts of firing during fictive locomotion. The data are consistent with a role for RCs in limiting the firing rates of motoneurons and IaINs during each burst.     

Deletions of rhythmic motoneuron activity during fictive locomotion and scratch provide clues to the organization of the mammalian central pattern generator

We examined the features of spontaneous deletions of bursts of motoneuron activity that can occur within otherwise rhythmic alternating flexor and extensor activity during fictive locomotion and scratch in adult decerebrate cats. Deletions of activity were observed both in hindlimb flexor and extensor motoneuron pools during brain stem-stimulation-evoked fictive locomotion but only in extensors during fictive scratch. Paired intracellular motoneuron recordings showed that deletions reduced the depolarization of homonymous motoneurons in qualitatively similar ways. Differences occurred in the extent to which activity in synergist motoneuron pools operating at other joints within the limb was reduced during deletions. The timing of the rhythmic activity that followed a deletion was often at an integer multiple of the preexisting locomotor or scratch cycle period. This maintenance of cycle period was also seen during deletions in which there was a complete failure of motoneuron depolarization. The activity of antagonist motoneurons was usually sustained during deletions with some rhythmic modulation at intervals of the preexisting cycle period. We discuss an organization of the central pattern generator for locomotion and scratch that functions as a single rhythm generator with separate and multiple pattern formation modules for controlling the hyper- and depolarization of subsets of motoneurons within the limb.     

Stumbling corrective reaction during fictive locomotion in the cat

An obstacle contacting the dorsal surface of a cat's hind foot during the swing phase of locomotion evokes a reflex (the stumbling corrective reaction) that lifts the foot and extends the ankle to avoid falling. We show that the same sequence of ipsilateral hindlimb motoneuron activity can be evoked in decerebrate cats during fictive locomotion. As recorded in the peripheral nerves, twice threshold intensity stimulation of the cutaneous superficial peroneal (SP) nerve during the flexion phase produced a very brief excitation of ankle flexors (e.g., tibialis anterior and peroneus longus) that was followed by an inhibition for the duration of the stimulus train (10-25 shocks, 200 Hz). Extensor digitorum longus was always, and hip flexor (sartorius) activity was sometimes, inhibited during SP stimulation. At the same time, knee flexor and the normally quiescent ankle extensor motoneurons were recruited (mean latencies 4 and 16 ms) with SP stimulation during fictive stumbling correction. After the stimulus train, ankle extensor activity fell silent, and there was an excitation of hip, knee, and ankle flexors. The ongoing flexion phase was often prolonged. Hip extensors were also recruited in some fictive stumbling trials. Only the SP nerve was effective in evoking stumbling correction. Delivered during extension, SP stimulus trains increased ongoing extensor motoneuron activity as well as increasing ipsilateral hip, knee, and ankle hindlimb flexor activity in the subsequent step cycle. The fictive stumbling corrective reflex seems functionally similar to that evoked in intact, awake animals and involves a fixed pattern of short-latency reflexes as well as actions evoked through the lumbar circuitry responsible for the generation of rhythmic alternating locomotion.     

Activity of interneurons within the L4 spinal segment of the cat during brainstem-evoked fictive locomotion

Summary Extracellular recordings from interneurons located in the L4 spinal segment were made during fictive locomotion produced by electrical stimulation of the mesencephalic locomotor region (MLR) in the paralysed decerebrate cat. Only interneurons within the L4 segment which received group II input from quadriceps, sartorius or the pretibial flexor muscle afferents and which had axonal projections to motor nuclei in L7 were selected for analysis. During the fictive step cycle two thirds of these interneurons fired action potentials during the time of activity in the ipsilateral hindlimb flexor neurograms. These cells were also less responsive to peripheral input during the extension phase of the fictive locomotion cycle. The remaining one third of the interneurons examined were not rhythmically active during locomotion. The possible contributions of the midlumbar interneurons to motoneuron activity during locomotion are discussed.     

Motoneuron input-resistance changes during fictive locomotion produced by stimulation of the mesencephalic locomotor region

Input-resistance changes during fictive locomotion were monitored in a variety of extensor and flexor hindlimb alpha-motoneurons in precollicular, postmammillary decerebrate cats induced to "walk" by electrical stimulation of the mesencephalic locomotor region (MLR). Using intracellular recording techniques and injected hyperpolarizing current pulses, the changes in the motoneuron input resistance recorded at the motoneuron soma were examined during nonlocomoting control periods as well as during the depolarized and hyperpolarized phases of the membrane potential oscillations (locomotor drive potentials, or LDPs) of fictive locomotion. In 28 of the 52 motoneurons examined, no change in the input resistance between the control and locomotor periods was observed. The remainder of the cells displayed a decrease (less than 20%) in input resistance when fictive stepping commenced. Over 80% of all the motoneurons depolarized (mean depolarization 4 mV), whereas only one LG motoneuron hyperpolarized (2 mV) with the onset of stimulation of the MLR. The remaining motoneurons did not display such changes. In 43 out of 52 motoneurons examined, no significant change in the input resistance could be observed between the depolarized and hyperpolarized phases of the step cycle. A decrease in the input resistance during the depolarized phase of the LDP was observed in four LG motoneurons, whereas five other motoneurons (2 LG, 1 TA, 1 PB, and 1 ST) displayed an increased input resistance during the depolarized phase compared with the hyperpolarized phase of locomotion. The data are consistent with the presence of an excitatory synaptic input alternating with an inhibitory input to the motoneuron during the fictive step cycle.(ABSTRACT TRUNCATED AT 250 WORDS)     

Modulation of short latency cutaneous excitation in flexor and extensor motoneurons during fictive locomotion in the cat

Summary We examined modulation of transmission in short-latency, distal hindlimb cutaneous reflex pathways during fictive locomotion in 19 decerebrate cats. Fictive stepping was produced either by electrical stimulation of the mesencephalic locomotor region (MLR) or by administration of Nialamide and 1-DOPA to acutely spinalized animals. Postsynaptic potentials (PSPs) produced by electrical stimulation of low threshold afferents (< 2.5 times threshold) in the superficial peroneal (SP), sural, saphenous or medial plantar nerves were recorded intracellularly from various extensor (n = 28) and flexor (n = 24) motoneurons and averaged throughout the step cycle, together with voltage responses to intrasomatic constant current pulses (in order to monitor relative cell input resistance). Each motoneuron studied displayed rhythmic background oscillations in membrane potential and correlated variations in input resistance. The average input resistance of extensor motoneurons was lowest during mid-flexion, when the cells were relatively hyperpolarized and silent. Conversely, average input resistance of flexor motoneurons was highest during mid-flexion, when they were depolarized and active. The amplitude of the minimum-latency excitatory components of PSPs produced by cutaneous nerve stimulation were measured from computer averaged records representing six subdivisions of the fictive step cycle. Oligosynaptic EPSP components were consistently modulated only in the superficial peroneal responses in flexor motoneurons, which exhibited enhanced amplitude during the flexion phase. With the other skin nerves tested (sural, saphenous, and plantar), no consistent patterns of modulation were observed during fictive locomotion. We conclude that transmission through some, but not all, oligosynaptic excitatory cutaneous pathways is enhanced by premotoneuronal mechanisms during the flexion phase of fictive stepping in several cat hindlimb motor nuclei. The present results suggest that the patterns of interaction between the locomotor central pattern generator and excitatory cutaneous reflex pathways depend on the source of afferent input and on the identity of the target motoneuron population.     

Intracellular analysis of reflex pathways underlying the stumbling corrective reaction during fictive locomotion in the cat

In cat and humans, contact between an obstacle and the dorsum of the foot evokes the stumbling corrective reaction (reflex) that lifts the foot to avoid falling. This reflex can also be evoked by short trains of stimuli to the cutaneous superficial peroneal (SP) nerve in decerebrate cats during the flexion phase of fictive locomotion. Here we examine intracellular events in hindlimb motoneurons accompanying stumbling correction. SP stimulation delivered during the flexion phase excites knee flexor motoneurons at short latency [minimum excitatory postsynaptic potential (EPSP) latency 1.8 ms; mean 2.7 ms]. Although a similar short latency excitation occurs in ankle extensors (mean latency, 2.8 ms), recruitment is delayed until successive shocks in the stimulus train overcome the locomotor-related hyperpolarization of ankle extensors. In ankle flexor motoneurons, SP stimulation evokes an inhibition (mean latency, 2.7 ms) that briefly reduces or stops their firing during the flexion phase. There is a phase-dependent modulation of SP-evoked EPSP amplitude as well as latency during locomotion. However, the more obvious change in SP reflex pathways with the onset of fictive locomotion is the reduced inhibition of ankle extensor motoneurons and the increased inhibition of ankle flexors. These results show that the characteristic pattern of hindlimb motoneuron activation during SP nerve-evoked stumbling correction results from 1) di- and trisynaptic excitation of knee flexor and ankle extensor motoneurons; 2) increased inhibitory postsynaptic potentials in ankle flexors and a suppression of inhibition in extensors, 3) sculpting of the short-latency SP postsynaptic effects by motoneuron membrane potential, and 4) longer latency excitatory effects that are likely evoked by lumbar interneurons involved in the generation of fictive locomotion.     

Mechanism for activation of locomotor centers in the spinal cord by stimulation of the mesencephalic locomotor region

The synaptic pathways of mesencephalic locomotor region (MLR)-evoked excitatory and inhibitory postsynaptic potentials (EPSPs and IPSPs) recorded from lumbar motoneurons of unanesthetized decerebrate cats during fictive locomotion were analyzed prior to, during, and after cold block of the medial reticular formation (MedRF) or the low thoracic ventral funiculus (VF). As others have shown, electrical stimulation of the MLR typically evoked short-latency excitatory or mixed excitatory/inhibitory PSPs in flexor and extensor motoneurons. The bulbospinal conduction velocities averaged approximately 88 m/s (range: 62-145 m/s) and segmental latencies for EPSPs ranged from 1.2 to 10.9 ms. The histogram of segmental latencies showed three peaks, suggesting di-, tri-, and polysynaptic linkages. Segmental latencies for IPSPs suggested trisynaptic or polysynaptic transmission. Most EPSPs (69/77) were significantly larger during the depolarized phase of the intracellular locomotor drive potential (LDP), and most IPSPs (35/46) were larger during the corresponding hyperpolarized phase. Bilateral cooling of the MedRF reversibly abolished locomotion of both hindlimbs as measured from the electroneurogram (ENG) activity of muscle nerves and simultaneously abolished or diminished the motoneuron PSPs and LDPs. Unilateral cooling of the VF blocked locomotion ipsilaterally and diminished it contralaterally with concomitant loss or decrease the motoneuron PSPs and LDPs. Relative to the side of motoneuron recording, cooling of the ipsilateral VF sometimes uncovered longer-latency EPSPs, whereas cooling of the contralateral VF abolished longer-latency EPSPs. It is concluded that MLR stimulation activates a pathway that relays in the MedRF and descends bilaterally in the VF to contact spinal interneurons that project to motoneurons. Local segmental pathways that activate or inhibit motoneurons during MLR-evoked fictive locomotion appear to be both ipsilateral and contralateral.     

Modelling spinal circuitry involved in locomotor pattern generation: insights from the effects of afferent stimulation

A computational model of the mammalian spinal cord circuitry incorporating a two-level central pattern generator (CPG) with separate half-centre rhythm generator (RG) and pattern formation (PF) networks has been developed from observations obtained during fictive locomotion in decerebrate cats. Sensory afferents have been incorporated in the model to study the effects of afferent stimulation on locomotor phase switching and step cycle period and on the firing patterns of flexor and extensor motoneurones. Here we show that this CPG structure can be integrated with reflex circuits to reproduce the reorganization of group I reflex pathways occurring during locomotion. During the extensor phase of fictive locomotion, activation of extensor muscle group I afferents increases extensor motoneurone activity and prolongs the extensor phase. This extensor phase prolongation may occur with or without a resetting of the locomotor cycle, which (according to the model) depends on the degree to which sensory input affects the RG and PF circuits, respectively. The same stimulation delivered during flexion produces a temporary resetting to extension without changing the timing of following locomotor cycles. The model reproduces this behaviour by suggesting that this sensory input influences the PF network without affecting the RG. The model also suggests that the different effects of flexor muscle nerve afferent stimulation observed experimentally (phase prolongation versus resetting) result from opposing influences of flexor group I and II afferents on the PF and RG circuits controlling the activity of flexor and extensor motoneurones. The results of modelling provide insights into proprioceptive control of locomotion.     

Toe flexor muscle spindle discharge and stretch modulation during locomotor activity in the decerebrate cat

In order to investigate the nature (i.e. static or dynamic) of fusimotor drive to the flexor hallucis longus (FHL) and flexor digitorum longus (FDL) muscles during locomotion we recorded Ia and group II muscle spindle afferent responses to sinusoidal stretch (0.25 and 1 mm amplitude, respectively, 4–5 Hz) in a decerebrate cat preparation. FHL Ia and group II afferents generally had increased discharge rates and decreased modulation to stretch throughout the step cycle, compared to rest, suggesting raised static γ drive at all locomotor phases. Although the modulation of Ia afferents was reduced during locomotion, most (13 of 18) showed a clear increasing trend during homonymous muscle activity (extension). This was consistent with phasic dynamic γ drive to FHL spindles linked with α drive. In agreement with previous reports, FHL gave a single burst of EMG activity during the step cycle while FDL α drive had two components. One was related to extension while the other comprised a brief burst around the end of this phase. Typically FDL Ia and group II afferents also had elevated firing rates and reduced modulation at all locomotor phases, again implicating static γ drive. Half the afferents (seven Ia, three group II) showed increased discharge during extension, suggesting phasic static γ drive. There was no γ drive associated with the late FDL α burst. In conclusion, the γ drives to FHL and FDL differed during locomotion. FHL, which has the α drive of a classic extensor, received γ drive that closely resembled other extensors. The γ drive of FDL, which exhibits both extensor and flexor α synergies, did not match either muscle type. These observations are compatible with the view that fusimotor drive varies in different muscles during locomotion according to the prevailing sensorimotor requirements.     

Excitability changes of ankle extensor group Ia and Ib fibers during fictive locomotion in the cat

Summary The present study examines the modulation of gastrocnemius-soleus (GS) monosynaptic reflexes as well as the intraspinal threshold changes of GS group I primary afferent terminals ending in the intermediate and motor nuclei during fictive locomotion in high decerebrate cats. The amplitude of the monosynaptic reflexes (MSR's) evoked in the medial gastrocnemius by stimulation of the lateral gastrocnemius nerve was increased during the extensor (E) phase, decreased during the flexion (F) phase of the step cycle and remainded transiently increased after spontaneous episodes of fictive stepping. The intraspinal threshold of populations and of single group Ia GS afferent fibers ending in the motor pool, as well as of single Ia and Ib fibers ending in the intermediate nucleus, showed a sustained reduction during the episodes of fictive locomotion with superimposed cyclic changes in phase with the step cycle. During fictive walking and trotting the reduction of the intraspinal threshold of both Ia and Ib fiber terminals was maximal during the middle or late portion of the F-phase. During fictive gallop elicited by stimulation of the superficial peroneus nerve, the decrease in the intraspinal threshold of the Ia afferent fibers occurred however in phase with the activity of the GS motoneurons. During episodes of fictive locomotion slow, sustained negative DC potential shifts lasting tents of seconds, reflecting an increase in the extracellular potassium concentration were recorded at the base of the dorsal horn and in the intermediate nucleus. The present findings support the existence of tonic and phasic depolarization of the intraspinal terminals of GS group Ia and Ib primary afferents during spontaneous fictive locomotion. It is suggested that accumulation of potassium ions in the extracellular space contributes mainly to the sustained depolarization of group I fibers. The phasic depolarization would be mostly due to the activation of specific sets of interneurons and may, in the case of Ia fibers, contribute to the cyclic modulation of the MRS elicited during fictive locomotion.     

Parallel reflex pathways from flexor muscle afferents evoking resetting and flexion enhancement during fictive locomotion and scratch in the cat

Reflex actions of muscle afferents in hindlimb flexor nerves were examined on ipsilateral motoneurone activity recorded in peripheral nerves during midbrain stimulation-evoked fictive locomotion and during fictive scratch in decerebrate cats. Trains of stimuli (15–30 shocks at 200 Hz) were delivered during the flexion phase at intensities sufficient to activate both group I and II afferents (5 times threshold, T ). In many preparations tibialis anterior (TA) nerve stimulation terminated ongoing flexion and reset the locomotor cycle to extension (19/31 experiments) while extensor digitorum longus (EDL) stimulation increased and prolonged the ongoing flexor phase activity (20/33 preparations). The effects of sartorius, iliopsoas and peroneus longus muscle afferent stimulation were qualitatively similar to those of EDL nerve. Resetting to extension was seen only with higher intensity stimulation (5 T ) while ongoing flexor activity was often enhanced at group I intensity (2 T ) stimulation. The effects of flexor nerve stimulation were qualitatively similar during fictive scratch. Reflex reversals were consistently observed in some fictive locomotor preparations. In those cases, EDL stimulation produced a resetting to extension and TA stimulation prolonged the ongoing flexion phase. Occasionally reflex reversals occurred spontaneously during only one of several stimulus presentations. The variable and opposite actions of flexor afferents on the locomotor step cycle indicate the existence of parallel spinal reflex pathways. A hypothetical organization of reflex pathways from flexor muscle afferents to the spinal pattern generator networks with competing actions of group I and group II afferents on the flexor and extensor portions of this central circuitry is proposed.     

Disynaptic excitation from the medial longitudinal fasciculus to lumbosacral motoneurons: modulation by repetitive activation,descending pathways,and locomotion

Summary Short-latency excitatory postsynaptic potentials (EPSPs) evoked by stimulation in the medial longitudinal fasciculus (MLF) were recorded intracellularly from motoneurons in the cat lumbosacral spinal cord. Monosynaptic and disynaptic EPSPs occurred in most flexor and extensor motoneurons studied. These EPSPs resulted from the activation of fast (> 100 m/s) descending axons from the MLF to the spinal cord. Several features distinguished monosynaptic and disynaptic MLF EPSPs. Disynaptic EPSPs exhibited temporal facilitation during short trains of stimulation, whereas monosynaptic EPSPs did not. Disynaptic EPSPs, but not monosynaptic EPSPs, were also facilitated by stimulation of the pyramidal tract and the mesencephalic locomotor region. However, disynaptic MLF EPSPs exhibited little or no facilitation when conditioned by short-latency cutaneous pathways. During fictive locomotion, the amplitude of disynaptic MLF EPSPs was modulated, with maximal amplitudes during the step cycle phase when the recorded motoneuron was active, resulting in reciprocal patterns of modulation of flexors and extensors. No comparable change was seen in the amplitude of monosynaptic MLF EPSPs during fictive stepping. These data suggest that the central pattern generator for locomotion modulates disynaptic MLF excitation at a premotoneuronal level in a phase-dependent manner. The effects of lesions made in the MLF and thoracic cord suggest that the interneurons in the disynaptic pathway from the MLF to motoneurons reside in the lumbosacral cord.     

Phasic control of the transmission in the excitatory and inhibitory reflex pathways from cutaneous afferents to α-motoneurones during fictive locomotion in cats

In high spinal paralyzed cats the effect of cutaneous nerve stimulation on lumbar motoneurons was investigated during fictive locomotion. EPSPs evoked from the cutaneous afferents were generally larger during the active phase of the motoneurones, while IPSPs tended to increase during the reciprocal phase. In some cases EPSPs occurred during the active phase, while IPSPs dominated during the reciprocal phase. Apparently, the transmission in the excitatory and inhibitory segmental reflex pathways from cutaneous afferents to α-motoneurones depends on the phase of the step cycle, but there is no general phase dependent alternating switching between these two pathways.     

Phasic modulation of short latency cutaneous excitation in flexor digitorum longus motoneurons during fictive locomotion

Summary We examined modulation of transmission of short-latency excitation produced by distal hindlimb cutaneous input, as well as fluctuations in motoneuron membrane potential and input resistance, in flexor digitorum longus (FDL) motoneurons during fictive locomotion. Fictive stepping was induced in unaesthetized, decerebrate cats either by repetitive stimulation of the mesencephalic locomotor region (MLR) or by administration of Nialamide and 1 DOPA after low spinal section. In the MLR preparations, brief depolarizing waves occurred in FDL cells during the early flexion phase of fictive stepping, immediately after cessation of activity in extensor muscles. In some FDL cells, plateau-like depolarizations also occurred during the extensor phase. Fictive stepping induced in acutely spinalized cats by administration of l-DOPA was slower and more variable; peak polarization in FDL motoneurons always occurred during the early flexion phase but there was usually no distinct depolarization during extension. In both types of preparation, the initial EPSP components in synaptic potentials (SP-EPSPs) produced by electrical stimulation of the cutaneous division of the superficial peroneal nerve (SP) were maximally facilitated during early flexion, coincident with the peak of background depolarization. This enhancement was manifested by an increase in the amplitude of initial SP-EPSP components or by decreased central latency of the initial EPSP components, or both. In most FDL motoneurons, input resistance decreased systematically during late flexion, coincident with relative membrane hyperpolarization. Correction of SP-EPSP amplitudes for changes in input resistance suggested that SP-EPSP facilitation persisted throughout the flexion phase These findings are discussed with reference to modulation of cutaneous reflexes during locomotion and the possibility that excitatory last-order interneurons in particular cutaneous reflex pathways may distribute excitatory drive from the central pattern generator for locomotion to FDL -motoneurons     

Proprioceptive input resets central locomotor rhythm in the spinal cat

Summary The reflex regulation of stepping is an important factor in adapting the step cycle to changes in the environment. The present experiments have examined the influence of muscle proprioceptors on centrally generated rhythmic locomotor activity in decerebrate unanesthetized cats with a spinal transection at Th12. Fictive locomotion, recorded as alternating activity in hindlimb flexor and extensor nerves, was induced by administration of nialamide (a monoamine oxidase inhibitor) and L-DOPA. Brief electrical stimulation of group I afferents from knee and ankle extensors were effective in resetting fictive locomotion in a coordinated fashion. An extensor group I volley delivered during a flexor burst would abruptly terminate the flexor activity and initiate an extensor burst. The same stimulus given during an extensor burst prolonged the extensor activity while delaying the appearance of the following flexor burst. Intracellular recordings from motoneurones revealed that these actions were mediated at premotoneuronal levels resulting from a distribution of inhibition to centres generating flexor bursts and excitation of centres generating extensor bursts. These results indicate that extensor group I afferents have access to central rhythm generators and suggest that this may be of importance in the reflex regulation of stepping. Experiments utilizing natural stimulation of muscle receptors demonstrate that the group I input to the rhythm generators arises mainly from Golgi tendon organ Ib afferents. Thus an increased load of limb extensors during the stance phase would enhance and prolong extensor activity while simultaneously delaying the transition to the swing phase of the step cycle.     

Activity of renshaw cells during fictive scratch reflex in the cat

Summary The experiments were performed on decerebrate curarized cats with a hindlimb either completely deafferented or partly deefferented. Through tactile stimulation of the pinna, a fictive scratch reflex was evoked and activity in muscle efferents was then observed, similar to that in actual scratching. The duration of the cycle was about 250 ms, with extensor activity during a short period of about 50 ms (S phase) and flexor activity during a much longer one (about 200 ms; L phase). Appearance of rhythmic bursts of discharges was preceded by tonic flexor activity (tonic phase of scratching). Discharges of Renshaw cells were recorded extracellularly during these three phases, in parallel with discharges in the gastrochemius-soleus nerve. During the tonic phase of the scratch reflex, some Renshaw cells with input from flexors decreased their activity while others increased. No change in Renshaw cell activity with input from extensors was then observed. During the rhythmic phases of the scratch reflex a majority of Renshaw cells was phasically active. They usually responded once per cycle, with a burst of 1–30 impulses of 50–100 ms duration, most often occurring at the end of the L phase and during the S phase. Bursts of Renshaw cells with input from flexors and of extensors, respectively, overlapped to a high degree. However, maximal firing of extensor-coupled Renshaw cells occurred somewhat later than that of flexor-coupled cells. Flexor-coupled Renshaw cells discharged mainly at the end of the L phase and during the S phase, i.e. when the flexor moto-neurones terminated their activity. Extensor-coupled Renshaw cells reached maximal activity during the S phase, i.e. when the extensor motoneurones were recruited. After spinal transection at C1 level, Renshaw cells responded with an increased number of spikes but without change in timing of the discharges during the scratch cycle. Most of the contralaterally located Renshaw cells studied were also phasically active during the scratch reflex. The role of motoneurones and spinal interneurones in determining the timing of Renshaw cell activity and the role of the latter in control of posture and rhythmic movements are discussed.     

Analysis of Ia-inhibitory synaptic input to cat spinal motoneurons evoked by vibration of antagonist muscles.

1. Steady-state inhibitory postsynaptic potentials (IPSPs) were evoked in tibialis anterior and extensor digitorum longus motoneurons of the cat by using tendon vibration to activate Ia-afferent fibers from the antagonist medial gastrocnemius muscle. 2. The effective synaptic currents (IN) underlying the steady-state IPSPs were measured by the use of a modified voltage-clamp technique. The amplitudes of the effective synaptic currents (1.62 +/- 0.66 nA, mean +/- SD; n = 20) extended over a fivefold range (0.5-2.7 nA) but were not correlated with the intrinsic properties of the motoneurons or with putative motor unit type. 3. We calculated the synaptic conductance (GS) underlying the steady-state Ia IPSPs from measurements of motoneuron input conductance during the activation of the Ia synaptic input. As was expected from Ohm's law, the Ia-inhibitory GS and IN were correlated (r = 0.49; P less than 0.05). Like IN, GS (175 +/- 202 nS, mean +/- SD; n = 20) was not correlated with the intrinsic properties of the motoneurons. 4. As has been reported previously for transient Ia IPSPs, the amplitudes of the steady-state IPSPs were correlated with motoneuron input resistance (r = 0.74; P less than 0.001) and homonymous Ia excitatory postsynaptic synaptic potential (EPSP) amplitude (r = 0.72; P less than 0.001). 5. The amplitudes of the steady-state Ia IPSPs and the homonymous Ia EPSPs were plotted on logarithmic axes. The slope (0.59) was significantly less than 1, which indicates that the gradient of Ia inhibition across the motoneuron pool is less steep than that of Ia excitation.(ABSTRACT TRUNCATED AT 250 WORDS)     

Locomotor activities in the decerebrate bird without phasic afferent input

We examined whether forelimb and hindlimb phasic afferent input is a prerequisite for the production of avian locomotor patterns. We eliminated phasic afferent feedback through paralysis of a decerebrate animal. The term "fictive" has been used to describe the neural activity associated with spontaneous or evoked motor output during neuromuscular paralysis. We observed that a paralysed decerebrate bird is capable of producing similar locomotor activity patterns as an unparalysed preparation, regardless of whether the "fictive" locomotion is generated spontaneously, or in response to focal electrical and/or neurochemical stimulation of discrete brainstem locomotor regions. Not all aspects of "fictive" locomotor patterns were identical to the locomotion elicited prior to paralysis. The stimulus current threshold necessary to evoke hindlimb locomotion increased from 69 +/- 22 mu A (mean +/- S.D.) prior to paralysis to 185 +/- 87 mu A for "fictive" stepping. For wing activity, the threshold increased from 84 +/- 46 mu A during wing flapping to 228 +/- 148 mu A for "fictive" flight. In addition, the frequency of "fictive" efferent locomotor activity from the leg nerve (1.04 +/- 0.44 Hz) decreased relative to the frequency of leg activity prior to paralysis (1.55 +/- 0.70 Hz). Similarly, the frequency of wing activity decreased from 2.73 +/- 0.73 Hz before paralysis to 1.8 +/- 0.69 Hz after paralysis. Finally flexor burst duration remained constant during treadmill and "fictive" walking while the extensor burst duration was markedly increased during "fictive" walking. Thus, the relative contributions of leg flexor activity to the overall step cycle (burst proportion = burst duration/cycle duration) decreased during evoked "fictive" stepping, while the burst proportion of the leg extensor increased. Afferent feedback therefore appears to modulate leg extensor burst duration more than leg flexor duration. For the wings, the burst proportion of the major wing depressors remained constant before and after paralysis.     

Distribution of oligosynaptic group I input to the cat medial gastrocnemius motoneuron pool

To characterize the oligosynaptic group I afferent input to the cat medial gastrocneumius (MG) motoneuron pool, the medial branch of the tibial nerve (MTIB: flexor digitorum and hallucis longus, popliteus, tibialis posterior and interosseous nerves), the nerves to flexor digitorum and hallucis longus (FDHL), or the nerves to the quadriceps muscles (QUAD) were stimulated at submaximal group I strength while recording intracellularly from MG motoneurons. Since previous work indicates that stimulation of these nerves at group I strength produces no significant monosynaptic Ia excitation or Renshaw inhibition of MG motoneurons, group I effects were assumed to be predominantly, though not exclusively, due to the action of Ib-fibers. Evidence supporting this assumption is presented in the following paper. MTIB, FDHL, and QUAD postsynaptic potentials (PSPs) were most commonly inhibitory. Since the MTIB, FDHL, and QUAD nerves are composed predominantly of fibers innervating muscles with extensor action, their inhibitory effect on MG motoneurons is consistent with previous findings that stimulation of Ib-afferents in nerves to extensor muscles produces di- and trisynaptic inhibition of extensor motoneurons. However, excitatory effects were observed in about one third of the motoneurons, indicating that oligosynaptic group I input is not homogeneously distributed within the MG motoneuron pool. Variations in QUAD, FDHL, and MTIB PSP pattern and amplitude were correlated with variations in the PSP pattern evoked by stimulation of the sural nerve: excitatory oligosynaptic group I PSPs generally appeared in motoneurons receiving excitatory cutaneous (sural nerve) input, whereas inhibitory PSPs generally appeared in motoneurons receiving some inhibitory cutaneous input and were largest in motoneurons receiving predominantly inhibition from the sural nerve. These variations in QUAD, FDHL, and MTIB PSP pattern and amplitude were not due to variations in resting potential and were only partly due to variations in intrinsic motoneuron properties or motoneuron "type." Our results indicate that activation of these cutaneous and group I muscle afferents can exert similar effects on the MG motoneuron pool. Moreover, the presence of a strong correlation between the distributions of cutaneous and oligosynaptic group I PSPs within a single motoneuron pool is consistent with the results of previous studies that have shown that some of the input to motoneurons from these peripheral afferents is mediated through common interneurons.