1. Recordings were made of local field potential (slow waves) and pyramidal tract neurone (PTN) discharge from pairs of sites separated by a horizontal distance of up to 1·5 mm in the primary motor cortex of two conscious macaque monkeys performing a precision grip task. 2. In both monkeys, the slow wave recordings showed bursts of oscillations in the 20-30 Hz range. Spectral analysis revealed that the oscillations were coherent between the two simultaneously recorded cortical sites. In the monkey from which most data were recorded, the mean frequency of peak coherence was 23·4 Hz. 3. Coherence in this frequency range was also seen between cortical slow wave recordings and rectified EMG of hand and forearm muscles active during the task, and between pairs of rectified EMGs. 4. The dynamics of the coherence were investigated by analysing short, quasi-stationary data segments aligned relative to task performance. This revealed that the 20-30 Hz coherent oscillations were present mainly during the hold phase of the precision grip task. 5. The spikes of identified PTNs were used to compile spike-triggered averages of the slow wave recordings. Oscillations were seen in 11Ï17 averages of the slow wave recorded on the same electrode as the triggering spike, and 11Ï17 averages of the slow wave recorded on the distant electrode. The mean period of these oscillations was 45·8 ms. 6. It is concluded that oscillations in the range 20-30 Hz are present in monkey motor cortex, are coherent between spatially separated cortical sites, and encompass the pyramidal tract output neurones. They are discernable in the EMG of active muscles, and show a consistent task-dependent modulation.5816

Field potential recordings from motor cortex show oscillations in the beta-band (∼20 Hz), which are coherent with similar oscillations in the activity of contralateral contracting muscles. Recent findings have revised concepts of how this activity might be generated in the cortex, suggesting it could achieve useful computation. Other evidence shows that these oscillations engage not just motor structures, but also return from muscle to the central nervous system via feedback afferent pathways. Somatosensory cortex has strong beta-band oscillations, which are synchronised with those in motor cortex, allowing oscillatory sensory reafference to be interpreted in the context of the oscillatory motor command which produced it.

Cortical oscillations have been the target of many recent investigations, because it has been proposed that they could function to solve the "binding" problem. In the motor cortex, oscillatory activity has been reported at a variety of frequencies between approximately 4 and approximately 60 Hz. Previous research has shown that 15-30 Hz oscillatory activity in the primary motor cortex is coherent or phase locked to activity in contralateral hand and forearm muscles during isometric contractions. However, the function of this oscillatory activity remains unclear. Is it simply an epiphenomenon or is it related to specific motor parameters? In this study, we investigated task-dependent modulation in coherence between motor cortex and hand muscles during precision grip tasks. Twelve right-handed subjects used index finger and thumb to grip two levers that were under robotic control. Each lever was fitted with a sensitive force gauge. Subjects received visual feedback of lever force levels and were instructed to keep them within target boxes throughout each trial. Surface EMGs were recorded from four hand and forearm muscles, and magnetoencephalography (MEG) was recorded using a 306 channel neuromagnetometer. All subjects showed significant levels of coherence (0.086-0.599) between MEG and muscle in the 15-30 Hz range. Coherence was significantly smaller when the task was performed under an isometric condition (levers fixed) compared with a compliant condition in which subjects moved the levers against a spring-like load. Furthermore, there was a positive, significant relationship between the level of coherence and the degree of lever compliance. These results argue in favor of coherence between cortex and muscle being related to specific parameters of hand motor function.

Damage to the corticospinal tract is a leading cause of motor disability, for example in stroke or spinal cord injury. Some function usually recovers, but whether plasticity of undamaged ipsilaterally descending corticospinal axons and/or brainstem pathways such as the reticulospinal tract contributes to recovery is unknown. Here, we examined the connectivity in these pathways to motor neurons after recovery from corticospinal lesions. Extensive unilateral lesions of the medullary corticospinal fibres in the pyramidal tract were made in three adult macaque monkeys. After an initial contralateral flaccid paralysis, motor function rapidly recovered, after which all animals were capable of climbing and supporting their weight by gripping the cage bars with the contralesional hand. In one animal where experimental testing was carried out, there was (as expected) no recovery of fine independent finger movements. Around 6 months post-lesion, intracellular recordings were made from 167 motor neurons innervating hand and forearm muscles. Synaptic responses evoked by stimulating the unlesioned ipsilateral pyramidal tract and the medial longitudinal fasciculus were recorded and compared with control responses in 207 motor neurons from six unlesioned animals. Input from the ipsilateral pyramidal tract was rare and weak in both lesioned and control animals, suggesting a limited role for this pathway in functional recovery. In contrast, mono- and disynaptic excitatory post-synaptic potentials elicited from the medial longitudinal fasciculus significantly increased in average size after recovery, but only in motor neurons innervating forearm flexor and intrinsic hand muscles, not in forearm extensor motor neurons. We conclude that reticulospinal systems sub-serve some of the functional recovery after corticospinal lesions. The imbalanced strengthening of connections to flexor, but not extensor, motor neurons mirrors the extensor weakness and flexor spasm which in neurological experience is a common limitation to recovery in stroke survivors.

The planning and execution of movements involve areas of the cerebral cortex that are separated in their activity both temporally and spatially. However, the mechanism by which these areas combine to produce a co-ordinated movement is unknown. Studies of the activity of neurones in the sensorimotor cortex in monkeys and humans have shown synchronous oscillatory activity in the 15-30 Hz range (Murthy & Fetz, 1992, 1996aSanes & Donoghue, 1993;Baker et al. 1997;Donoghue et al. 1998). These synchronous oscillations have been suggested to link the disparate motor signals together in a manner analogous to the perceptual 'binding' of stimulus attributes in the visual cortex (Singer & Gray, 1995), where the observed neuronal synchronization is attributed to neurones that participate in the encoding of related information. Sanes & Donoghue (1993) and Murthy & Fetz (1996a) have shown that oscillations in monkey sensorimotor cortex are synchronous over large distances (up to 14 mm), suggesting the involvement of large neuronal populations. However, Murthy & Fetz (1996b) concluded that the oscillatory episodes had no consistent relationship with a variety of motor tasks, and suggested that rather than being involved directly in 'binding' during movement execution, the oscillations could be a neuronal correlate of attention during sensorimotor tasks. 1. Recent reports have shown task-related changes in oscillatory activity in the 15-30 Hz range in the sensorimotor cortex of human subjects and monkeys during skilled hand movements. In the monkey these oscillations have been shown to be coherent with oscillatory activity in the electromyographic activity of hand and forearm muscles. 2. In this study we investigated the modulation of oscillations in the electromyogram (EMG) of human volunteers during tasks requiring precision grip of two spring-loaded levers. 3. Two tasks were investigated: in the 'hold' task, subjects were required to maintain a steady grip force (ca 2·1 N or 2·6 N) for 8 s. In the 'ramp' task, there was an initial hold period for 3 s (force ca 2·1 N) followed by a linear increase in grip force over a 2 s period. The task ended with a further steady hold for 3 s at the higher force level (ca 2·6 N). 4. Surface EMGs were recorded from five hand and forearm muscles in 12 subjects. The coherence of oscillatory activity was calculated between each muscle pair. Frequencies between 1 and 100 Hz were analysed. 5. Each subject showed a peak in the coherence spectra in the 15-30 Hz bandwidth during the hold task. This coherence was absent during the initial movement of the levers. During the ramp task the coherence in the 15-30 Hz range was also significantly reduced during the movement phase, and significantly increased during the second hold period, relative to the initial hold. 6. There was coherence between the simultaneously recorded magnetoencephalogram (MEG) and EMG during steady grip in the hold task; this coherence disappeared during the initial lever movement. Using a single equivalent current dipole source...

The primate reticulospinal tract is usually considered to control proximal and axial muscles, and to be involved mainly in gross movements such as locomotion, reaching and posture. This contrasts with the corticospinal tract, which is thought to be involved in fine control, particularly of independent finger movements. Recent data provide evidence that the reticulospinal tract can exert some influence over hand movements. Although clearly secondary to the corticospinal tract in healthy function, this could assume considerable importance after corticospinal lesion (such as following stroke), when reticulospinal systems could provide a substrate for some recovery of function. We need to understand more about the abilities of the reticular formation to process sensory input and guide motor output, so that rehabilitation strategies can be optimised to work with the innate capabilities of reticular motor control.

There is currently much interest in the synchronisation of neural discharge and the potential role it may play in information coding within the nervous system. We describe some recent results from investigations of synchronisation within the motor system. Local field potentials (LFPs) and identified pyramidal tract neurones (PTNs) were recorded from the primary motor cortex of monkeys trained to perform a precision grip task. The LFPs showed bursts of oscillatory activity at 20-30 Hz, which were coherent with the rectified electromyographs (EMG) of contralateral hand and forearm muscles. This oscillatory synchronisation showed a highly specific task dependence, being present only during the part of the task when the animal maintained a steady grip and not during the movement phases before or after it. PTNs were phase-locked to LFP oscillations, implying that at least part of the coherence between cortical activity and EMG was mediated by corticospinal fibres. The phase locking of the PTNs to LFP oscillations produced task-dependent oscillatory synchronisation between PTN pairs, as assessed by the single-unit cross-correlation histogram. Recordings were also made from normal human subjects performing a precision grip similar to that used in the monkey recordings. Pairs of EMGs recorded from intrinsic hand and forearm muscles showed 20-30 Hz coherence, which modulated during task performance, being present only during periods of steady contraction. We suggest that these changes in EMG-EMG synchronisation reflect changing levels of synchronous drive from the corticospinal system. The generation of oscillations in the cortex is discussed in the light of results from a model of local cortical circuits. Other modelling work has shown that synchrony in the corticospinal inputs could act to recruit motoneurones more efficiently, producing more output force from a muscle than asynchronous inputs firing at the same mean rate. A speculative hypothesis is presented on the role of synchronous oscillations in the motor system, which is consistent with experimental observations to date.

Although a major descending motor pathway in mammals, the reticulospinal tract’s contribution to upper limb control in primates has received relatively little attention. Reticulospinal connections are widely assumed to be responsible for coordinated gross movements primarily of proximal muscles, whereas the corticospinal tract mediates fine movements, particularly of the hand. In this study, we employed intracellular recording in anaesthetised monkeys to examine the synaptic connections between the reticulospinal tract and antidromically identified cervical ventral horn motoneurons, focussing in particular on motoneurons projecting distally to wrist and digit muscles. We found that motoneurons receive mono- and disynaptic reticulospinal inputs, including monosynaptic excitatory connections to motoneurons that innervate intrinsic hand muscles, a connection not previously known to exist. We show that excitatory reticulomotoneuronal connections are as common and as strong in hand motoneuron groups as in forearm or upper arm motoneurons. These data suggest that the primate reticulospinal system may form a parallel pathway to distal muscles, alongside the corticospinal tract. Reticulospinal neurons are therefore in a position to influence upper limb muscle activity after damage to the corticospinal system as may occur in stroke or spinal cord injury, and may be a target site for therapeutic interventions.