Neuromuscular patterns of stereotypic hindlimb behaviors in the first two postnatal months. II. Stepping in spinal kittens

From birth to postnatal day 60, neuromuscular patterns for airstepping and treadmill stepping were assessed in kittens spinalized (T12) at birth (Day-1) or at the end of the second postnatal week (Day-14). Within 72 h after spinalization, all kittens displayed stepping motions, but exteroceptive facilitation (e.g. tail pinch) was required to initiate and sustain both behaviors. In Day-14 spinal kittens, the hindlimbs spontaneously and alternately airstepped, but in Day-1 spinal kittens exteroceptive stimulation was usually necessary to evoke airstepping, and the hindlimbs stepped synchronously. Kittens in both groups developed atypical neuromuscular patterns; flexor bursts were nearly twice as long in duration as extensor bursts. Development of bipedal treadmill stepping was similar for Day-1 and Day-14 spinal kittens, but differed from that for normal kittens. Tested at the same belt speeds, stepping was more easily elicited in spinal kittens, bouts of repetitive stepping were longer, and cycle periods were shorter than in normal kittens until postnatal week 6. Spinal kittens, however, seldom exhibited adequate weight support during hindlimbs stepping, and the neuromuscular patterns associated with bipedal stepping were atypical. For spinal kittens, the relationship between the extensor burst duration and the cycle period was reduced substantially, and flexor activity was initiated earlier in the step cycle and was longer in duration than that for normal kittens. These atypical intralimb synergies may have been the consequence of altered lumbosacral circuits produced by the spinal transection. It is also possible that these spinal circuits, lacking rostral input, were particularly susceptible to abnormal motion-dependent feedback resulting from reduced hindlimb weight support.     

Neuromuscular patterns of stereotypic hindlimb behaviors in the first two postnatal months. I. Stepping in normal kittens

Neuromuscular patterns associated with the development of hindlimb stepping behaviors were studied from birth to postnatal day 60 in normal kittens. Hindlimb muscles were chronically implanted with EMG electrodes at birth to characterize interlimb coordination and intralimb synergies during development of overground and treadmill stepping. Airstepping was also examined but seldom occurred after the second postnatal week. All kittens performed stepping under each condition, including weight-supported stepping, by postnatal day 3. The number of sequential steps on the treadmill and overground increased with age and cycle periods decreased. At onset, stepping behaviors were characterized by adult-like EMG patterns. Interlimb coordination was typified by alternating extensor bursts of similar duration. Extensors at the knee and ankle were coactive during the stance phase, and extensor burst durations were strongly correlated with the cycle periods over a wide range of stepping frequency. Ankle flexor and extensor muscles were reciprocally active during postural tremor, bouts of airstepping, and weight-supported steps on the treadmill and overground. The duration of the reciprocal flexor bust did not vary with cycle period or age. Observations of stepping behaviors and adult-like EMG patterns during initial postnatal development were contingent on optimal testing conditions. Taken together, the data suggest that pattern-generating circuits for regulating interlimb coordination and intralimb muscle synergies are potentially functional prior to the normal ontogenetic onset of locomotion. Perhaps the prolonged postnatal development of locomotion reflects the time required to establish adaptive mechanisms, such as postural control and agility, rather than spinal pattern-generating circuits for locomotion.     

Early onset of hindlimb paw-shake responses in spinal kittens: new perspective in motor development

Normal kittens and kittens spinalized (T12) at 14 days of age were tested for onset of paw-shake responses (PSR) in fore and hindlimbs. In normals, onset followed a cephalocaudal pattern that was coincident with the development of stable posture, 21-28 days of age. In spinals, onset of hindlimb PSRs preceded that of the forelimb and occurred soon after cordotomy, independent of posture development. These findings suggest that in the neonate, spinal networks responsible for coordinating PSRs are normally inhibited until the response can be supported by stable posture.     

Electromyographic activity patterns of ankle flexor and extensor muscles during spontaneous and L-DOPA-induced locomotion in freely moving neonatal rats

In rats, hindlimb postural and locomotor functions mature during the first 3 postnatal weeks. Previous evidence indicates that maturation of descending monoaminergic pathways is important for the postnatal emergence of locomotion with adequate antigravity postural support. Here we have studied the effect of the monoamine precursor L-DOPA on locomotor activity in freely moving postnatal rats (7-9 days old) using electromyographic recordings from ankle extensor (soleus) and flexor (tibialis anterior or extensor digitorum longus) muscles. Before pharmacological treatment, both muscles were usually silent at rest, and during spontaneous movements there was a high degree of coactivation between the two antagonists. This was due to a longer electromyographic (EMG) burst duration in flexors, which partly overlapped with the extensor burst. L-DOPA administration (150 mg/kg) resulted in a marked increase in postural tonic EMG activity in extensors which appeared gradually within 10 min after injection and was sufficient for the pups to maintain a standing posture with the pelvis raised above ground. Thereafter, episodes of locomotion characterized by rhythmic reciprocal bursts of EMG activity in flexor and extensor muscles were seen. The L-DOPA-induced rhythmic EMG pattern was also seen in postnatal rats subjected to a midthoracic spinal cord transection, indicating that the effect of L-DOPA on motor coordination is exerted primarily at the level of the spinal pattern generator. Analysis of EMG burst characteristics showed that the pattern of L-DOPA-induced locomotion in both intact and spinalized postnatal rats resembled in some respects that observed in adults during spontaneous locomotion. The appearance of reciprocal activation during L-DOPA-induced locomotion in neonates was primarily due to a shortening of the EMG burst duration in flexors, which reduced the degree of antagonist coactivation. These results show that the spinal cord has the potential to produce coordinated overground locomotion several days before such movements are normally expressed in the freely moving animal.     

Motor pattern deletions and modular organization of turtle spinal cord

The turtle spinal cord contains a central pattern generator (CPG) that produces rhythmic hindlimb motor patterns during a rostral scratch. This review describes evidence in support of the hypothesis that the turtle rostral scratch CPG has a modular structure similar to that described in the Unit-Burst-Generator hypothesis for cat locomotion by Grillner. During normal rostral scratch in turtle, activity bursts rhythmically alternate with quiescence for each motor neuron pool; agonist activity rhythmically alternates with antagonist activity at each degree of freedom, e.g., hip, knee; and a transition from knee flexor to knee extensor motor neuron activity occurs midway during each hip flexor motor neuron burst. Hip extensor deletions, knee flexor deletions, and knee extensor deletions are motor pattern variations of rostral scratch. During each of these variations, agonist activity is rhythmic; antagonist activity and agonist quiescence are absent. Several classes of evidence during both normal and variation motor patterns support a modular organization of the turtle rostral scratch CPG: electroneurographic recordings from axons of motor neurons, intracellular recordings of synaptic potentials in motor neurons, and extracellular unit recordings from spinal interneurons. These data support the hypotheses that the knee extensor module is different from the hip extensor module and that the knee flexor module is different from the hip flexor module. Potential mechanisms for rhythmogenesis include reciprocal connections between agonist and antagonist modules at each degree of freedom, and agonist module rhythmogenesis. Additional tests of the modular hypothesis for turtle rostral scratch include unit recordings from knee-related interneurons during normal rostral scratch, as well as during knee-related deletions.     

Development and characteristics of airstepping in chronic spinal cats

Airstepping, walking-like movements of the hindlimbs, is a commonly observed behavior in chronic spinal animals when they are held vertically. The purpose of this study was to: describe the development of airstepping after spinalization and compare it to the onset of segmental reflexes, characterize the EMG pattern of muscle activity during spontaneous airstepping, and examine the effects of sensory perturbation on the characteristic pattern. Airstepping was analyzed during three conditions of tonic sensory perturbation which included: tail pinching, tape applied to one hindpaw, and immobilization of the ankle and knee in a plaster cast. Seven adult cats were spinalized at T-12, and bipolar electrode wires were surgically implanted in selected hindlimb muscles at the hip, knee, and ankle. Testing began within 48 hr of transection. Segmental reflexes and paw-shake responses were present in the first week; however, the earliest observed airstepping occurred during tail pinching at 2 weeks after surgery, and the average onset of spontaneous airstepping (without exteroceptive stimuli) was at 33 days. The average cycle period of spontaneous airstepping (691 msec) was comparable to the shortest periods reported for fictive rhythms and to treadmill walking between 1 and 2 m/sec. Intralimb coordination was characterized by flexor and extensor synergies typical of locomotion, while interlimb coordination was characterized by alternating cycles similar to that reported for treadmill walking and fictive locomotion. Neither intralimb nor interlimb patterns of coordination were altered by conditions of sensory perturbation, although cycle period and EMG recruitment level were variable. Many characteristics of airstepping are similar to those of treadmill and fictive locomotion.(ABSTRACT TRUNCATED AT 250 WORDS)     

Weight-bearing hindlimb stepping in treadmill-exercised adult spinal cats

Hindlimb locomotion on a motor-driven treadmill was studied in 5 cats spinalized at a low thoracic level adults. Six months after surgery, the cats were anesthetized and implanted for electromyographic (EMG) and force recordings in hindlimb muscles. For the last 5 months of the spinalization period, the hindlimbs of each cat were exercised daily for 30 minutes on a treadmill. Data were collected during hindlimb locomotion on a treadmill across the entire range of speeds each cat could accommodate. All trials were filmed (100 frames/s) for kinematic analysis. EMG data were recorded from the soleus (Sol), medial gastrocnemius (MG), tibialis anterior (TA) and extensor digitorum longus (EDL). Forces were recorded in vivo from the Sol and MG tendons. All cats could sustain full weight-bearing stepping without the need for mechanical stimulation of the tail. Although the general stepping pattern of the spinal cats was remarkably similar to that of normal cats, several key differences were identified. Compared to normal cats, the adult spinal cats walked at a lower range of speeds and exhibited a longer swing phase duration. The Sol produced forces and displayed activation periods comparable to those observed in normal cats. The MG of adult spinal cats, however, produced lower forces and had a later onset of activation in comparison to normal cats. Each of the muscles in all spinal cats exhibited tremor during stepping. These results suggest that there were limitations in the activation levels of some hindlimb flexor and extensor muscles during treadmill locomotion. These data further suggest that, in normal cats, accommodation to treadmill speed is accomplished by modulating supraspinal input to the lumbar spinal cord while leaving many of the timing details to be regulated by lumbar spinal networks.     

Messages conveyed by spinocerebellar pathways during scratching in the cat. II. Activity of neurons of the ventral spinocerebellar tract

(1) Signals transmitted to the cerebellum by the spino-reticulocerebellar pathway (SRCP) during scratching were studied. For this purpose, the activity of neurons of the lateral reticular nucleus (LRN), which are the last-order neurons of the SRCP, was recorded during scratching in thalamic cats. Scratching was evoked by stimulation of the pinna. LRN neurons were identified antidromically by stimulation of the hindlimb area in the cerebellar anterior lobe. In most experiments, animals were immobilized with Flaxedil, and stimulation of the pinna resulted in fictitious scratching, i.e., in periodical reciprocal activity of flexor and extensor motoneurons typical of actual scratching. (2) During both actual and fictitious scratching, the discharge frequency of LRN neurons was rhythmically modulated in relation with the scratch cycle. Most LRN neurons fired in short high-frequency bursts of spikes which coincided (completely or partly) with the extensor phase of the cycle. In this respect the SRCP differs from the ventral spinocerebellar tract (VSCT) which is maximally active in the flexor phase of the cycle. (3) The firing pattern of LRN neurons during fictitious scratching was similar to that during actual scratching. Therefore, the rhythmical burst firing of LRN neurons is determined mainly by the central mechanisms and not by the rhythmical sensory input. (4) Rhythmical modulation of LRN neurons disappeared after transection of the ipsilateral lateral funiculus of the spinal cord in which spinoreticular fibers are located. On the other hand, considerable reduction of rhythmical activity in descending brainstem-spinal pathways after contralateral hemisection of the spinal cord did not affect the discharge pattern of LRN neurons. These two facts indicate that the SRCP conveys mainly messages about activity of the central spinal mechanisms, and that influences of supraspinal motor centers.on LRN neurons and on spinoreticular neurons are of minor importance. (5) Axonal terminations of LRN neurons are distributed rather evenly over the hindlimb area in the anterior lobe of the cerebellum. Therefore, messages about the events, which happen within the spinal cord in the vicinity of the extensor phase of the cycle, arrive at every point of the hindlimb area.     

Spinalization unmasks clonidine's alpha 1-adrenergic mediated excitation of the flexor reflex in rats

Clonidine exerts alpha 2-adrenergic mediated depressant effects on most behaviors measured in a normal animal. However, in the spinal-transected (spinalized) animal, clonidine apparently facilitates the flexor reflex through a stimulation of spinal alpha 1-adrenoceptors. The purpose of the present study was to determine if spinalization per se causes the shift in clonidine's profile from an alpha 2- to an alpha 1-adrenergic agonist. The hindlimb flexor reflex was elicited by electrical pulses delivered through electrodes implanted subcutaneously in the hindpaw and was measured with a force transducer and polygraph. In contrast to an alpha 2-adrenergic mediated inhibition of the flexor reflex in intact rats, clonidine produced an alpha 1-adrenergic mediated increase in flexor reflex amplitude in spinalized rats. Because decerebration did not alter the depression due to clonidine, and intraventricular (but not intrathecal) administration of oxymetazoline mimicked the effect of clonidine, the depressant effects of alpha 2-adrenergic agonists are mediated through alpha 2-adrenergic receptors localized in the brainstem. Alternate methods for inducing a functional spinal transection (spinal block with intrathecal procaine; spinal ligation) indicated that the shift in clonidine's effect from inhibition of the flexor reflex to excitation occurred immediately following spinalization. Spinal ligation did not produce alpha 1-adrenergic supersensitivity at 15 min or 2 hr after transection, as measured by alterations in [3H]prazosin receptor binding or behavioral responses to clonidine. Thus, the shift in clonidine's effects from alpha 2-adrenergic mediated inhibition of the flexor reflex in intact rats to alpha 1-adrenergic mediated excitation in spinalized rats results because spinal transection unmasks clonidine's alpha 1-adrenergic stimulatory effect. Other conditions under which clonidine exerts alpha 1-adrenoceptor mediated excitatory effects on behavior are discussed.     

Stepping behaviors in chronic spinal cats with one hindlimb deafferented

Adult cats develop spontaneous airstepping (walking motions without ground contact) 4-6 weeks after spinal transection (Giuliani and Smith, 1985). This unique preparation provides an in vivo model for studying the effects of hindlimb deafferentation on stepping behaviors without rostral input to lumbosacral segments. The primary purpose of this study was to characterize airstepping and, to a lesser extent, bipedal treadmill locomotion in chronic spinal cats after deafferentation. Five cats were spinalized at T12, and EMG electrodes were implanted in selected knee and ankle muscles. After spontaneous airstepping developed, one hindlimb was deafferented extensively. Movements of both hindlimbs were depressed following unilateral deafferentation. By the third week, spontaneous airstepping recovered in the normal hindlimb (N-Limb), but in the deafferented limb (D-Limb) airstepping was not spontaneous during the 3-4 months of testing. By the seventh week, bilateral airstepping was facilitated by tonic exteroceptive simulation (tail-pinch). During bouts of bilateral airstepping, assessed during the third month of deafferentation, D-Limb airstep cycles were characterized by erratic rhythm. Further, cycle periods and extensor burst durations were prolonged, but flexor burst durations were unmodified. In contrast, N-Limb cycles were rhythmic and of normal duration, but extensor burst durations were short and unrelated to cycle periods. Interlimb coordination was markedly unstable, showing only brief periods of alternating steps. When erratic bilateral airstepping was accompanied by micturition, rhythmic and alternate stepping emerged, with normalized intralimb synergies. During hindlimb treadmill locomotion, tested in 2 cats, the N-Limb took weight-bearing steps and followed changes in the belt speed. The D-Limb, however, stepped erratically, making contact on the dorsum of the paw; consistent bouts by bilateral stepping were not obtained. Lumbosacral afference of some type, either from the hindlimb or from regions remote from the limb, such as the bladder, appears to be essential for stabilizing the coupling between hindlimb generators for stable interlimb coordination. Future modeling of interlimb coordination should consider the role of afference.     

Different effects of spinalization and locomotor training of spinal animals on cholinergic innervation of the soleus and tibialis anterior motoneurons

Cholinergic input modulates excitability of motoneurons and plays an important role in the control of locomotion in both intact and spinalized animals. However, spinal cord transection in adult rats affects cholinergic innervation of only some hindlimb motoneurons, suggesting that specificity of this response is related to functional differences between motoneurons. Our aim was therefore to compare cholinergic input to motoneurons innervating the soleus (Sol) and tibialis anterior (TA) motoneurons following spinal cord transection at a low-thoracic level. The second aim was to investigate whether deficits in cholinergic input to these motoneurons could be modified by locomotor training. The Sol and TA motoneurons were identified by retrograde labelling with fluorescent dyes injected intramuscularly. Cholinergic terminals were detected using anti-vesicular acetylcholine transporter (VAChT) antibody. Overall innervation of motoneurons was evaluated with anti-synaptophysin antibody. After spinalization we found a decrease in the number of VAChT-positive boutons apposing perikarya of the Sol (to 49%) but not TA motoneurons. Locomotor training, resulting in moderate functional improvement, partly reduced the deficit in cholinergic innervation of Sol motoneurons by increasing the number of VAChT-positive boutons. However, the optical density of VAChT-positive boutons terminating on various motoneurons, which decreased after spinalization, continued to decrease despite the training, suggesting an impairment of acetylcholine availability in the terminals. Different effects of spinal cord transection on cholinergic innervation of motoneurons controlling the ankle extensor and flexor muscles point to different functional states of these muscles in paraplegia as a possible source of activity-dependent signaling regulating cholinergic input to the motoneurons.     

Changes in the efferent activity of the scratching generator evoked by external mechanical force applied to a hindlimb in the decerebrate cat]

Changes in the afferent inflow appeared with application of the external mechanical force to the rhythmically moving hindlimb have been studied for their scratching generator activity in decerebrate adult cats. Phasic relationship between violent hindlimb transference and efferent activity of the scratching generator was opposite to that found earlier during fictitious scratching, which is explained by difference between afferent activity during real scratching movement and passive movement of the hindlimb in the immobilized animal. Maximum redistribution in the relationship between flexor and extensor intensities in response to the stimulation corresponds to maximum rate of the violent hindlimb transference.     

Activity of rubrospinal neurons during locomotion and scratching in the cat

It is now well established that locomotion and scratching in vertebrates can result from the activation of a spinal central generator. The possibility of control of these rhythmic motor activities by the red nucleus has been analyzed in the thalamic cat, in which efferent nerve discharges representing fictive locomotion or fictive scratching can still be recorded following paralysis by curarization. It was found that the discharge of lumbar-projecting rubrospinal neurons is modulated in relation to the intensity and frequency of the rhythmic efferent activity in the contralateral hindlimb. The average firing frequency was minimal at the transition between the extensor and flexor efferent bursts and increased progressively to reach a maximum in the second part of the flexor burst. Comparison of the rubrospinal activities during real and fictive rhythmic motor activities revealed only minor influences of phasic afferent inputs. Analysis of the relations between the rhythmic discharges found in rubrospinal neurons, cerebellar neurons (interpositus nucleus and paravermal Purkinje cells of the cerebellar anterior lobe) and neurons of an ascending pathway (ventral spinocerebellar tract) leads to the conclusion that the rubrospinal tract belongs to an internal loop between spinal and supraspinal centres. However, until now, the results do not allow the evaluation of its contribution to the motor performance, even in situations which, like those studied here, do not involve the complex motor control present in the intact cat.     

Changes in the excitability of motor neurons and synaptic effects on them during formation of the generator of scratching movements in the cat

It is shown that the state of the segmental apparatus of the lumbal spinal cord of immobilized intercollicularly decerebrated cats during formation of a scratching generator is essentially different from that of a spinal animal. The excitability of "aiming" and "scratching" motoneurons increases, recurrent and reciprocal Ia inhibition of motoneurons becomes stronger and influence of Ib afferents weaker. The flexor reflex afferents exert inhibitory influences on "aiming" motoneurons. After spinalization these influences become excitatory, and the inhibitory influences on "scratching" motoneurons become weaker. The functional role of described changes is discussed.     

Development of Locomotor Behavior in the Spinal Kitten

This study was undertaken to determine the locomotor capability of kittens whose spinal cords were transected at birth. The postnatal development of reflex and goal-directed locomotion was examined during the first 5 postnatal months in kittens that received low thoracic spinal cord transections as newborns. Some spinal kittens developed aberrant quadrupedal forms of locomotion. The onset of quadrupedal locomotion, however, was delayed by 2-3 months compared to the normal kitten (42) and deteriorated by 5 months of age. Qualitative and quantitative analyses demonstrated that the quadrupedal locomotion was abnormal. Although some step cycles were characterized by full weight support, the typical hindlimb step cycle of the best performing cat showed inadequate weight support and balance. No spinal cat was able to coordinate the hindlimbs with the forelimbs during overground locomotion on a runaway or during quadrupedal locomotion on a treadmill. Neuroanatomical tracing with WGA-HRP and immunocytochemical techniques showed no axonal regeneration or growth into or across the lesion sites. The aberrant form of quadrupedal locomotion developed without descending input to the caudal spinal cord. The variability in performance among animals suggested that compensatory strategies were important factors in the spinal kitten's achievement of quadrupedal locomotion. Hindlimb weight-supported stepping during quadrupedal locomotion in some animals underscored the capacity of the isolated caudal spinal cord to generate both rhythmical stepping movements and weight support. The maintenance of developmentally immature, but functional, hindlimb postures suggested that the development of the isolated caudal spinal cord was arrested in the absence of descending input.     

Electromyographic Activity Patterns of Ankle Flexor and Extensor Muscles during Spontaneous and -DOPA-Induced Locomotion in Freely Moving Neonatal Rats

In rats, hindlimb postural and locomotor functions mature during the first 3 postnatal weeks. Previous evidence indicates that maturation of descending monoaminergic pathways is important for the postnatal emergence of locomotion with adequate antigravity postural support. Here we have studied the effect of the monoamine precursor -DOPA on locomotor activity in freely moving postnatal rats (7–9 days old) using electromyographic recordings from ankle extensor (soleus) and flexor (tibialis anterior or extensor digitorum longus) muscles. Before pharmacological treatment, both muscles were usually silent at rest, and during spontaneous movements there was a high degree of coactivation between the two antagonists. This was due to a longer electromyographic (EMG) burst duration in flexors, which partly overlapped with the extensor burst. -DOPA administration (150 mg/kg) resulted in a marked increase in postural tonic EMG activity in extensors which appeared gradually within 10 min after injection and was sufficient for the pups to maintain a standing posture with the pelvis raised above ground. Thereafter, episodes of locomotion characterized by rhythmic reciprocal bursts of EMG activity in flexor and extensor muscles were seen. The -DOPA-induced rhythmic EMG pattern was also seen in postnatal rats subjected to a midthoracic spinal cord transection, indicating that the effect of -DOPA on motor coordination is exerted primarily at the level of the spinal pattern generator. Analysis of EMG burst characteristics showed that the pattern of -DOPA-induced locomotion in both intact and spinalized postnatal rats resembled in some respects that observed in adults during spontaneous locomotion. The appearance of reciprocal activation during - DOPA-induced locomotion in neonates was primarily due to a shortening of the EMG burst duration in flexors, which reduced the degree of antagonist coactivation. These results show that the spinal cord has the potential to produce coordinated overground locomotion several days before such movements are normally expressed in the freely moving animal.     

Is spinal cord isolation a good model of muscle disuse?

The patterns of normal daily activity that are required to maintain normal skeletal muscle properties remain unknown. The present study was designed to determine whether spinal cord isolation can be used as a reliable experimental model of neuromuscular inactivity, that is, as a baseline for the absence of activity. Electromyograms (EMGs) were recorded from selected hindlimb muscles of unanesthetized rats over 24-hour periods before and 7, 30, 60, and 90 days after surgical isolation of the lumbar spinal cord. Our data indicate that some rat slow muscle fibers pre-surgery were activated for less than 3 hours per day. Spinal cord isolation (SI) reduced the mean daily integrated EMG (IEMG) and daily EMG duration in the primary slow extensor muscle (soleus) to <1% of control, and in the primary fast extensor muscles [medial gastrocnemius (MG) and vastus lateralis (VL)] to <2% of control. These parameters were decreased to <8% and 3% of control, respectively, in a primary fast flexor muscle, the tibialis anterior (TA). From 30 to 90 days post-SI, the mean amplitudes of the spontaneous EMG bursts were relatively normal in the soleus, increased approximately 2-fold in the MG and VL, and increased approximately 4-fold in the TA. Some evidence of the normal antagonistic flexor-extensor relationship was apparent in the brief periods of recorded activity post-SI. These results indicate that SI eliminates nearly all of the normal EMG activity in the hindlimb muscles in the presence of relatively normal muscle innervation and functional intraspinal neural circuitry.     

Characterization of synaptic potentials in hindlimb extensor motoneurons during L-DOPA-induced fictive locomotion in acute and chronic spinal cats

Intracellular recordings were carried out in extensor hindlimb motoneurons during L-DOPA-induced fictive locomotion from acute and chronically spinalized cats. It was demonstrated that motoneurons in chronic animals exhibited irregularly occurring, high frequency discharges within a given burst of a spike train, and sporadic membrane potential depolarizations and hyperpolarizations correlated with extensor and flexor nerve filament activity, respectively, during the locomotor cycle. In contrast, motoneurons recorded from acute cats demonstrated smooth membrane potential fluctuation and regularly occurring low frequency spike discharges. These results indicate that the pharmacologically activated central pattern generator (CPG) for locomotion in young adult chronic cats spinalized at two weeks of age produces disorganized locomotor-like patterns in the absence of sensory feedback. It is suggested that the above-mentioned characteristics of membrane potential fluctuations and spike discharges are not due to alterations in the motoneuron membrane properties, but instead are due to changes in the inputs to the motoneurons.     

Influence of spinalization on spinal withdrawal reflex responses varies depending on the submodality of the test stimulus and the experimental pathophysiological condition in the rat

The influence of midthoracic spinalization on thermally and mechanically induced spinal withdrawal reflex responses was studied in the rat. There were three experimental groups of rats: healthy controls, rats with a spinal nerve ligation-induced unilateral neuropathy, and rats with a carrageenan-induced inflammation of one hindpaw. Tail flick response was induced by radiant heat. Hindlimb withdrawal was induced by radiant heat, ice water, and innocuous or noxious mechanical stimulation of the paw. Prior to spinalization, spinal nerve ligated and carrageenan-treated animals had a marked unilateral allodynia and hyperalgesia. Spinalization tended to induce a facilitation of noxious heat-evoked reflexes. This spinalization-induced facilitation was stronger on tail than hindlimb withdrawal. Spinalization-induced skin temperature change did not explain the facilitation of noxious heat-evoked reflexes. In contrast, spinal withdrawal responses induced by noxious cold or mechanical stimulation were significantly suppressed following spinalization. The spinalization-induced facilitatory effects as well as inhibitory ones on spinal reflexes were enhanced in inflamed/neuropathic animals. The results indicate that the tonic descending control of spinal nocifensive responses varies depending on the submodality of the test stimulus, the segmental level of the reflex (tail vs. hindlimb), and on the pathophysiological condition.     

Characterization of synaptic potentials in hindlimb extensor motoneurons duringl-DOPA-induced fictive locomotion in acure and chronic spinal cats

Intracellular recordings were carried out in extensor hindlimb motoneurons duringl-DOPA-induced fictive locomotion from acute and chronically spinalized cats. It was demonstrated that motoneurons in chronic animals exhibited irregularly occuring, high frequency discharges within a given burst of a spike train, and sporadic membrane potential depolarizations and hyperpolarizations correlated with extensor and flexor nerve filament activity, respectively, during the locomotor cycle. In contrast, motoneurons recorded from acute cats demonstrated smooth membrane potential fluctuation and regularly occuring low frequency spike discharges. These results indicate that the pharmacologically activated central pattern generator (CPG) for locomotion in young adult chronic cats spinalized at two weeks of age produces disorganized locomotor-like patterns in the absence of sensory feedback. It is suggested that the above-mentioned characteristics of membrane potential fluctuations and spike discharges are not due to alterations in the motoneuron membrane properties, but instead are due to changes in the inputs to the motoneurons.