Modulation of coordinated muscle activity during imposed sinusoidal hip movements in human spinal cord injury

Individuals with chronic spinal cord injury (SCI) often demonstrate multijoint reflex activity that is clinically classified as an extensor spasm. These responses are commonly observed in conjunction with an imposed extension movement of the hips, such as movement from a sit to a supine position. Coincidentally, afferent feedback from hip proprioceptors has also been implicated in the control of locomotion in the spinalized cat. Because of this concurrence, we postulated that extensor spasms that are triggered by hip extension might involve activation of organized interneuronal circuits that also have a role in locomotion. If true, imposed oscillations of the hip would be expected to produce activity of the leg musculature in a locomotor pattern. Furthermore, this muscle activity would be entrained to the hip movement. The right hip joints of 10 individuals with chronic SCI, consisting of both complete [American Spinal Injury Association (ASIA) A] and incomplete (ASIA B,C) injuries, were subjected to ramp and hold (10 s) movements at 60 degrees /s and sinusoidal oscillations at 1.2, 1.88, and 2.2 rad/s over ranges from 40 to -15 degrees (+/-5 degrees ) using a custom servomotor system. Surface EMG from seven lower extremity muscles and sagittal-plane joint torques were recorded to characterize the response. Ramp and hold perturbations produced coactivation at the hip, knee, and ankle joints, with a long duration (5-10 s). Sinusoidal perturbations yielded consistent muscle timing patterns that resulted in alternating flexor and extensor joint torques. EMG and joint torques were commonly entrained to the frequency of movement, with rectus femoris, vastus medialis, and soleus activity coinciding with hip extension and medial hamstrings activity occurring during hip flexion. Individual muscle timing patterns were consistent with hip position during normal gait, except for the vastus medialis. These results suggest that reflexes associated with extensor spasms may occur through organized interneuronal pathways, such as spinal centers for locomotion.     

Motor unit firing during and after voluntary contractions of human thenar muscles weakened by spinal cord injury

Spinal cord injury may change both the distribution and the strength of the synaptic input within a motoneuron pool and therefore alter force gradation. Here, we have studied the relative contributions of motor unit recruitment and rate modulation to force gradation during voluntary contractions of thenar muscles performed by five individuals with chronic (>1 yr) cervical spinal cord injury. Mean +/- SD thenar unit firing rates were low during both steady-level 25% (8.3 +/- 2.2 Hz, n = 27 units) and 100% maximal voluntary contractions (MVCs, 9.2 +/- 3.1 Hz, n = 23 units). Thus modest rate modulation, or a lack of it in some units, was seen despite an average fourfold increase in integrated surface electromyographic activity and force. During ramp contractions, units were recruited at 5.7 +/- 2.5 Hz, but still only reached maximal firing rates of 12.8 +/- 4.9 Hz. Motor units were recruited up to 85% of the maximal force achieved (14.6 +/- 5.6 N). In contrast, unit recruitment in control hand muscles is largely complete by 30% MVC. Thus, during voluntary contractions of thenar muscles weakened by cervical spinal cord injury, motor unit rate modulation was limited and recruitment occurred over a wider than usual force range. Those motor units that were stopped voluntarily had significantly lower derecruitment versus recruitment thresholds. However, 8 units (24%) continued to fire long after the signal to end the voluntary contraction at a mean frequency of 5.9 +/- 0.8 Hz. The forces generated by this prolonged unit activity ranged from 0.3 to 7.2% maximum. Subjects were unable to stop this involuntary unit activity even with the help of feedback. The mechanisms that underlie this prolonged motor unit firing need to be explored further.     

Migration of motor pool activity in the spinal cord reflects body mechanics in human locomotion

During the evolution of bipedal modes of locomotion, a sequential rostrocaudal activation of trunk muscles due to the undulatory body movements was replaced by more complex and discrete bursts of activity. Nevertheless, the capacity for segmental rhythmogenesis and the rostrocaudal propagation of spinal cord activity has been conserved. In humans, motoneurons of different muscles are arranged in columns, with a specific grouping of muscles at any given segmental level. The muscle patterns of locomotor activity and the biomechanics of the body center of mass have been studied extensively, but their interrelationship remains poorly understood. Here we mapped the electromyographic activity recorded from 30 bilateral leg muscles onto the spinal cord in approximate rostrocaudal locations of the motoneuron pools during walking and running in humans. We found that the rostrocaudal displacements of the center of bilateral motoneuron activity mirrored the changes in the energy due to the center-of-body mass motion. The results suggest that biomechanical mechanisms of locomotion, such as the inverted pendulum in walking and the pogo-stick bouncing in running, may be tightly correlated with specific modes of progression of motor pool activity rostrocaudally in the spinal cord.     

Sustained muscle contractions maintained by autonomous neuronal activity within the human spinal cord

It is well known that muscle contraction can be easily evoked in the human soleus muscle by applying single-pulse electrical stimulation to the tibial nerve at the popliteal fossa. We herein reveal the unexpected phenomenon of muscle contractions that can be observed when train stimulation is used instead. We found, in 11 human subjects, that transient electrical train stimulation (1-ms pulses, 50 Hz, 2 s) was able to induce sustained muscle contractions in the soleus muscle that outlasted the stimulation period for greater than 1 min. Subjects were unaware of their own muscle activity, suggesting that this is an involuntary muscle contraction. In fact, the excitability of the primary motor cortex (M1) with the sustained muscle contractions evaluated by transcranial magnetic stimulation was lower than the excitability with voluntary muscle contractions even when both muscle contraction levels were matched. This finding indicates that M1 was less involved in maintaining the muscle contractions. Furthermore, the muscle contractions did not come from spontaneous activity of muscle fibers or from reverberating activity within closed neuronal circuits involving motoneurons. These conclusions were made based on the respective evidence: 1) the electromyographic activity was inhibited by stimulation of the common peroneal nerve that has inhibitory connections to the soleus motoneuron pool and 2) it was not abolished after stopping the reverberation (if any) for approximately 100 ms by inducing the silent period that followed an H-reflex. These findings indicate that the sustained muscle contractions induced in this study are most likely to be maintained by autonomous activity of motoneurons and/or interneurons within the human spinal cord.     

Five basic muscle activation patterns account for muscle activity during human locomotion

An electromyographic (EMG) activity pattern for individual muscles in the gait cycle exhibits a great deal of intersubject, intermuscle and context-dependent variability. Here we examined the issue of common underlying patterns by applying factor analysis to the set of EMG records obtained at different walking speeds and gravitational loads. To this end healthy subjects were asked to walk on a treadmill at speeds of 1, 2, 3 and 5kmh⁻¹ as well as when 35–95% of the body weight was supported using a harness. We recorded from 12–16 ipsilateral leg and trunk muscles using both surface and intramuscular recording and determined the average, normalized EMG of each record for 10–15 consecutive step cycles. We identified five basic underlying factors or component waveforms that can account for about 90% of the total waveform variance across different muscles during normal gait. Furthermore, while activation patterns of individual muscles could vary dramatically with speed and gravitational load, both the limb kinematics and the basic EMG components displayed only limited changes. Thus, we found a systematic phase shift of all five factors with speed in the same direction as the shift in the onset of the swing phase. This tendency for the factors to be timed according to the lift-off event supports the idea that the origin of the gait cycle generation is the propulsion rather than heel strike event. The basic invariance of the factors with walking speed and with body weight unloading implies that a few oscillating circuits drive the active muscles to produce the locomotion kinematics. A flexible and dynamic distribution of these basic components to the muscles may result from various descending and proprioceptive signals that depend on the kinematic and kinetic demands of the movements.     

Absence of local sign withdrawal in chronic human spinal cord injury

Local sign withdrawal, a reflex to direct the limb away from noxious cutaneous stimuli, is thought to be indicative of a modular organization of the spinal cord. To assess the integrity of such an organization of the spinal cord in chronic human spinal cord injury (SCI), we tested the electromyogram (EMG) and joint torque responses to cutaneous stimuli applied to 6 locations of the leg in 10 SCI volunteers and 3 spinal-intact controls. The 6 locations included the medial arch of the foot, the second metatarsal, the dorsum, the region over the sural nerve at the lateral malleolus, and the anterior and posterior aspects of the lower leg. Although spinal-intact subjects demonstrated local sign withdrawal, the data from SCI subjects indicated that an invariant flexion response pattern was produced regardless of stimulus location. Ankle dorsiflexion and hip flexion were produced in all subjects at all locations and no difference in the ratio of hip:ankle torques could be detected for the 6 test locations. A windup-crossover test, employing a sequence of 6 stimuli at 1-s intervals was used to assess whether common neuronal pathways were responsible for the loss of modular organization. An additional 10 SCI volunteers were tested using stimuli in which the stimulus location was switched between the 2nd and 3rd stimulus of the test sequence. The response to the crossover stimulus more closely resembled the response to the 3rd stimulus of a windup sequence than a response without conditioning stimuli. These results indicate that increased excitability produced by windup at one stimulus site is maintained at the 2nd site. This observation suggests that deep dorsal horn neurons, typically associated with musculotopic mapping, may be reorganized in chronic spinal cord injury.     

Phase-dependent electromyographic activity of the lower-limb muscles of a patient with clinically complete spinal cord injury during orthotic gait

We examined the lower-limb electromyographic (EMG) activity from a patient with clinically complete spinal cord injury during orthotic gait. A newly developed gait orthosis was used to obtain bipedal locomotion. The surface EMG data during the gait together with the biomechanical variables were collected by way of a radio EMG system. A cyclic EMG activation pattern corresponding to the gait cycles were observed in each of the paralyzed lower-limb muscles during the orthotic gait. Although the EMG activation did not seem to contribute toward generating the gait, it showed some similarities to that of the infant stepping or immature gait. These results might be regarded as one of the indirect pieces of evidence that suggest the existence of a spinally originating motor mechanism underlying human locomotion. Received: 5 September 1997 / Accepted: 1 December 1997     

Flexion reflex modulation during stepping in human spinal cord injury

The flexion reflex modulation pattern was investigated in nine people with a chronic spinal cord injury during stepping using body weight support on a treadmill and manual assistance by therapists. Body weight support was provided by an upper body harness and was adjusted for each subject to promote the best stepping pattern with the least manual assistance required by the therapists. The flexion reflex was elicited by sural nerve stimulation with a 30 ms pulse train at 1.2–2 times the tibialis anterior reflex threshold. During stepping, stimuli were randomly dispersed across the gait cycle which was divided into 16 equal bins. A long latency (>110 ms) flexion reflex was present in all subjects, while a short (>30 ms) and a medium latency (>70 ms) flexion reflex were present only in three subjects. For each response, the non-stimulated EMG was subtracted from the stimulated EMG at identical time windows and bins, normalized to the maximal corresponding EMG, and significant differences were established with a Wilcoxon rank-sum test. The long latency flexion reflex was facilitated at late stance and during the swing-to-stance transition phase. A reflex depression was present from heel strike until mid-stance and during the swing-to-stance transition phase. The short and medium latency flexion reflexes were depressed during mid-stance followed by facilitation during the stance-to-swing transition phase. Regardless of the latency, facilitatory flexion responses during the swing phase coincided with decreased activity of ipsilateral ankle extensors. The flexion reflex was modulated in a phase dependent manner, a behavior that was absent for the soleus H-reflex in most of these patients (Knikou et al. in Exp Brain Res 193:397–407, 2009). We propose that training should selectively target spinal reflex circuits in which extensor muscles and reflexes are involved in order to maximize sensorimotor recovery in these patients.     

Contribution of afferent feedback to the soleus muscle activity during human locomotion

During the stance phase of the human step cycle, the ankle undergoes a natural dorsiflexion that stretches the soleus muscle. The afferent feedback resulting from this stretch enhances the locomotor drive. In this study a robotic actuator was used to slightly enhance or reduce the natural ankle dorsiflexion, in essence, mimicking the small variations in the ankle dorsiflexion movement that take place during the stance phase of the step cycle. The soleus (SOL) and tibialis anterior EMG were analyzed in response to the ankle trajectory modifications. The dorsiflexion enhancements and reductions generated gradual increments and decrements, respectively, in the ongoing SOL EMG. We exercised care to ensure that the imposed ankle movements were too slow to elicit distinct burst-like stretch reflex responses that have been investigated previously. The increased SOL EMG after the dorsiflexion enhancements was reduced when the group Ia afferents were blocked with peripheral ischemia at the thigh, and during high-frequency Achilles tendon vibration. However, neither ischemia nor tendon vibration affected the decrements in the SOL EMG during the dorsiflexion reductions. These findings give evidence of the contribution of afferent feedback to the SOL activity in an ongoing basis during the stance phase. The results suggest that mainly feedback from the group Ia pathways is responsible for the increments in the SOL EMG during the dorsiflexion enhancements. However, the decrements in the SOL activity might be mediated by different afferent mechanisms.     

Increases in corticospinal tract function by treadmill training after incomplete spinal cord injury

In this study, we examined if several months of intensive locomotor training increases the function of spared corticospinal tract pathways after chronic spinal cord injury (SCI) in association with the recovery of locomotor function. Transcranial magnetic stimulation (TMS) at incrementing levels of intensity was applied over the motor cortex supplying either the tibialis anterior or vastus lateralis muscles, and the resulting peak-to-peak amplitude of the motor-evoked potentials (MEPs) were measured to obtain a recruitment curve both before and after training. In the majority of subjects (7/8), 3-5 mo of daily intensive training increased the responses to TMS in at least one of the leg muscles tested (9/13). On average, across all muscles tested MEP(max), which was evoked at high stimulation intensities, increased by 46% and MEP(h), which was evoked at intermediate stimulation intensities, increased by 45% (both significantly different from 0), indicating an increase in corticospinal tract connectivity from training. The slope of the sigmoid function fit to the recruitment curve increased by 24% after training (significantly different), indicating an expansion and/or increased excitability of corticospinal circuits supplying muscles to the lower leg. We also observed that the average duration of the silent period measured at MEP(max) increased after training from 130 to 178 ms, suggesting that training had effects on cortical circuits thought to mediate this long-lasting inhibition. The percentage increase in MEP(max) was positively and significantly correlated to the degree of locomotor recovery as assessed by the WISCI II score, the distance a subject could walk in 6 min, and the amplitude of the locomotor EMG activity, suggesting that the corticospinal tract, in part, mediated the functional locomotor recovery produced from training.     

Fatigue properties of human thenar motor units paralysed by chronic spinal cord injury

Human muscles paralysed chronically by spinal cord injury (SCI) fatigue excessively. Whether these reductions in force reflect a decrease in the fatigue resistance of the motor units is unknown. Our aim was to determine the fatigability of thenar motor units paralysed chronically (10 ± 2 years) by cervical SCI. Surface electromyographic activity (EMG) and force were recorded from 17 paralysed motor units ( n = 7 subjects) in response to intraneural motor axon stimulation (13 pulses at 40 Hz, 1 s⁻¹ for 2 min). Unit force decreased progressively, reaching 8–60% of initial after 2 min, whereas both the amplitude and area of the first EMG potentials in the trains increased significantly (both P < 0.05). Thus, transmission of neural signals to the sarcolemma was effective and the reduction in force must reflect impaired processes in the muscle fibres. The median fatigue index for paralysed units (0.31), the ratio of the force at 2 min compared to the initial force, was significantly lower than that for units from control subjects (0.85, P < 0.05), but the distribution of fatigue indices for each population had a similar shape (ranges: 0.08–0.60 and 0.41–0.95, respectively). Hence, chronic paralysis did not limit the range of fatigability typically found for thenar units, only its magnitude. These findings suggest that all paralysed units underwent similar reductions in fatigue resistance. After fatigue, paralysed unit forces were reduced at all frequencies (1–100 Hz, P < 0.05). Twitch contraction and half-relaxation times were increased, as was the frequency needed to produce half maximal force ( P < 0.05). Thus, stimulation protocols used to produce functional movements in paralysed muscles need to accommodate the significant and rapid fatigue of the motor units.     

Flexor reflex responses triggered by imposed knee extension in chronic human spinal cord injury

Hypersensitivity of the flexor reflex pathways to input from force-sensitive muscle afferents may contribute to the prevalence and severity of muscle spasms in patients with spinal cord injury (SCI). In this study, we triggered flexor reflexes with constant velocity knee movements in 15 subjects with SCI. Ramp and hold knee extension perturbations were imposed on one leg while the hip and ankle were held in an isometric position using an instrumented leg brace. Knee, ankle and hip torque responses and electromyograms from six muscles of the leg were recorded following controlled knee extension at four different velocities. Tests were conducted with the hip in both flexed and extended positions. During the movement into knee extension, a velocity-dependent stretch reflex, represented by a progressively increasing knee flexion torque, was observed. In addition, another type of reflex that resembled a flexor reflex (flexion of the hip and ankle) was also triggered by the imposed knee extension. The magnitude of the ankle dorsiflexion torque responses was significantly correlated to the stretch reflex torque at the knee in 9 of the 15 subjects. We concluded that stretch reflexes initiate a muscle contraction that then can contribute to a flexor reflex response, possibly through muscle group III/IV afferent pathways. These results suggest that spasticity in SCI consists of a myriad of complex reflex responses that extend beyond stretch reflexes.     

The effects of human ankle muscle vibration on posture and balance during adaptive locomotion

This study investigated the contribution of ankle muscle proprioception to the control of dynamic stability and lower limb kinematics during adaptive locomotion, by using mechanical vibration to alter the muscle spindle output of individuals' stance limbs. It was hypothesised that muscle length information from the ankle of the stance limb provides information describing location as well as acceleration of the centre of mass (COM) with respect to the support foot during the swing phase of locomotion. Our prediction, based on this hypothesis was that ankle muscle vibration would cause changes to the position and acceleration of the COM and/or compensatory postural responses. Vibrators were attached to both the stance limb ankle plantarflexors (at the Achilles tendon) and the opposing dorsiflexor muscle group (over tibialis anterior). Participants were required to walk along a 9-m travel path and step over any obstacles placed in their way. There were three task conditions: (1) an obstacle (15 cm in height) was positioned at the midpoint of the walkway prior to the start of the trial, (2) the same obstacle was triggered to appear unexpectedly one step in front of the participant at the walkway midpoint and (3) the subjects' walking path remained clear. The participants' starting position was manipulated so that the first step over the obstacle (when present) was always performed with their right leg. For each obstacle condition participants experienced the following vibration conditions: no vibration, vibration of the left leg calf muscles or vibration of the anterior compartment muscles of the lower left leg. Vibration began one step before the obstacle at left leg heel contact and continued for 1 s. Vibrating the ankle muscles of the stance limb during the step over an obstacle resulted in significant changes to COM behaviour [measured as displacement, acceleration and position with respect to the centre of pressure (COP)] in both the medial/lateral (M/L) and anterior/posterior planes. There were also significant task-specific changes in stepping behaviour associated with COM control (measured as peak M/L acceleration, M/L foot displacement and COP position under the stance foot during the step over the obstacle). The results provide strong evidence that the primary endings of ankle muscle spindles play a significant role in the control of posture and balance during the swing phase of locomotion by providing information describing the movement of the body's COM with respect to the support foot. Our results also provide supporting evidence for the proposal that there are context-dependent changes in muscle spindle sensitivity during human locomotion.     

Facilitation of synaptic activity in the frog spinal cord produced by 4-aminopyridine

Reflex activity in the isolated frog spinal cord is increased after adding 4-aminopyridine (4-AP) to the bath. Depression of synaptic transmission by increased magnesium or EGTA is effectively antagonized by 4-AP. It is suggested that 4-AP facilitates synaptic transmission in the central nervous system by interacting with the calcium mechanism involved in transmitter release from presynaptic nerve terminals.     

EMG activity in pectoral and femoral muscles during spinal cord cooling in exercising pigeons

Adult domestic pigeons, with thermodes chronically implanted in the vertebral canal, were trained to walk on a treadmill. In the first series of experiments, EMG activity in a pectoral (M. pectoralis) and a femoral muscle (M. biceps femoris) was measured to determine if shivering could occur during exercise. When the spinal cord was cooled (36.2±0.5°C) during exercise (0.6 km/h), pectoral muscle EMG activity increased by 80%, while femoral muscle EMG activity did not change significantly. EMG activity remained unchanged during exercise in control experiments where the spinal cord was not cooled.In the second series of experiments, the spinal cord was first cooled (36.1±0.5°C) for 5 min in resting pigeons and then the treadmill was started. Spinal cord cooling during rest significantly increased pectoral muscle EMG activity but not that of the femoral muscle. Within 1 s after the onset of exercise, EMG activity in the pectoral muscle decreased by 74%.In both series of experiments, shivering was not induced in the femoral muscle. The level of pectoral muscle EMG activity stimulated by spinal cord cooling during exercise in the first series of experiments corresponded to the level to which EMG activity was reduced by exercise during spinal cord cooling in the second series of experiments (192% and 186% in relation to the post-cooling level, respectively).It is concluded that shivering can be induced in the pectoral muscle by spinal cord cooling during exercise in the pigeon. The level of shivering induced in the pectoral muscle is well balanced according to the degree of spinal cord cooling (thermal information) and the running speed of exercise (non-thermal information possibly from active muscles and joints).     

Role of endogenous release of norepinephrine in muscle spasms after chronic spinal cord injury

The recovery of persistent inward currents (PICs) and motoneuron excitability after chronic spinal cord transection is mediated, in part, by the development of supersensitivity to residual serotonin (5HT) below the lesion. The purpose of this paper is to investigate if, like 5HT, endogenous sources of norepinephrine (NE) facilitate motoneuron PICs after chronic spinal transection. Cutaneous-evoked reflex responses in tail muscles of awake chronic spinal rats were measured after increasing presynaptic release of NE by administration of amphetamine. An increase in long-lasting reflexes, known to be mediated by the calcium component of the PIC (CaPIC), was observed even at low doses (0.1-0.2 mg/kg) of amphetamine. These findings were repeated in a reduced S2 in vitro preparation, demonstrating that the increased long-lasting reflexes by amphetamine were neural. Under intracellular voltage clamp, amphetamine application led to a large facilitation of the motoneuron CaPIC. This indicates that the increases in long-lasting reflexes induced by amphetamine in the awake animal were, in part, due to actions directly on the motoneuron. Reflex responses in acutely spinal animals were facilitated by amphetamine similar to chronic animals but only at doses that were ten times greater than that required in chronic animals (0.2 mg/kg chronic vs. 2.0 mg/kg acute), pointing to a development of supersensitivity to endogenous NE in chronic animals. In summary, the increases in long-lasting reflexes and associated motoneuron CaPICs by amphetamine are likely due to an increased release of endogenous NE, which motoneurons become supersensitive to in the chronic stages of spinal cord injury.     

Intramuscular EMG from the hip flexor muscles during human locomotion

The purpose was to investigate the activation pattern of five major hip flexor muscles and its adaptation to changing speed and mode of progression. A total of 11 healthy subjects performed walking and running on a motor-driven treadmill at speeds ranging from 1.0 to 6.0 m s^?1. Intramuscular fine-wire electrodes were used to record myoelectric signals from the iliacus, psoas, sartorius, rectus femoris and tensor fascia latae muscles. The basic pattern, with respect to number of activation periods, remained the same irrespective of speed and mode of progression. However, differences in the relative duration and timing of onset of activation occurred between individual muscles. Over the speed range in walking, a progressively earlier onset was generally seen for the activation period related to hip flexion. Changes in EMG amplitude were measured in the iliacus and psoas muscles and showed a marked increase and difference between walking and running at speeds above 2.0 m s^?1. Thus, the alternating flexion–extension movements at the hip during locomotion appear to be governed by a rather fixed `neural program' which normally only needs minor modulations to accomplish the adjustments accompanying an increase in speed of progression as well as a change from walking to running.     

Twitch and tetanic properties of human thenar motor units paralyzed by chronic spinal cord injury

Little is known about how human motor units respond to chronic paralysis. Our aim was to record surface electromyographic (EMG) signals, twitch forces, and tetanic forces from paralyzed motor units in the thenar muscles of individuals (n = 12) with chronic (1.5-19 yr) cervical spinal cord injury (SCI). Each motor unit was activated by intraneural stimulation of its motor axon using single pulses and trains of pulses at frequencies between 5 and 100 Hz. Paralyzed motor units (n = 48) had small EMGs and weak tetanic forces (n = 32 units) but strong twitch forces, resulting in half-maximal force being achieved at a median of only 8 Hz. The distributions for cumulative twitch and tetanic forces also separated less for paralyzed units than for control units, indicating that increases in stimulation frequency made a smaller relative contribution to the total force output in paralyzed muscles. Paralysis also induced slowing of conduction velocities, twitch contraction times and EMG durations. However, the elevated ratios between the twitch and the tetanic forces, but not contractile speed, correlated significantly with the extent to which unit force summated in response to different frequencies of stimulation. Despite changes in the absolute values of many electrical and mechanical properties of paralyzed motor units, most of the distributions shifted uniformly relative to those of thenar units obtained from control subjects. Thus human thenar muscles paralyzed by SCI retain a population of motor units with heterogeneous contractile properties because chronic paralysis influenced all of the motor units similarly.     

大鼠脊髓发育及损伤后NIDD mRNA表达的实验研究

目的：探讨在大鼠发育过程中及急性脊髓损伤后脊髓组织中NIDD （nNOS-interacting DHHC domain-containing protein with dendritic mRNA）mRNA的表达变化及意义。方法：采用改良Allen＇s打击法，咬除T8-10椎板后，造成大鼠脊髓损伤模型，致伤量为10×10g·cm；借助实时荧光定量PCR、原位杂交与免疫荧光结合的方法，定量、定位研究发育过程中及脊髓损伤后早期大鼠脊髓组织中NIDD mRNA与nNOS mRNA表达的时间和空间分布特征。结果：大鼠发育过程中，胚胎16d的大鼠脊髓中可见NIDD mRNA的高表达，在生后1d，与nNOS共表达于尚未分化成熟的前角，在白质也见NIDD的阳性信号。成年后呈低表达；nNOS mRNA于生后1～3d出现表达高峰；脊髓损伤后NIDD mRNA表达明显增多，在8h到达高峰，分布于脊髓前角、中间带、中央管周围及后角nNOS阳性的神经元，7d恢复至正常水平；nNOS mRNA在损伤后8h达到高峰，1d降低至正常水平。而且，在脊髓损伤后NIDD mRNA与nNOS mRNA二者表达呈正相关。结论：胎鼠脊髓中，NIDD高表达于nNOS阳性细胞，提示其在脊髓组织的发育成熟过程中的作用可能与nNOS相关。脊髓损伤后脊髓组织中NIDD与nNOS表达增多，提示在脊髓损伤的病理过程中，NIDD可能通过调节nNOS的细胞亚定位及活性而发挥一定的生物学作用。