Biomedical subjects
Publications and source records attributed to T Drew.
This article reviews the recent literature concerning the role of visual information in the control of locomotion with an emphasis on the neurophysiological mechanisms that underlie visually triggered, voluntary, gait modifications. Data are presented to show how these gait modifications may be encoded by the motor cortex, and how they may interact with the basic locomotor rhythm.
1. This article presents the results from stimulation in 21 loci within the medullary reticular formation (MRF; between 0.5 and 2.5 mm from the midline) and in 5 loci in the medial longitudinal fasciculus (MLF) of four intact, unanesthetized cats during locomotion. Stimulus trains (11 pulses, 0.2-ms duration, 330 Hz, stimulus strength 35 microA) were applied at those loci in each track at which the most widespread effects in each of the four limbs were obtained with the cat at rest. Electromyograms were recorded from flexor and extensor muscles of each limb. 2. As previously reported, stimulation with the cat at rest generally evoked brief, short-latency, twitch responses in both flexor and extensor muscles of more than one limb. In contrast, stimulation during locomotion evoked a more complex pattern of activity in which responses were normally evoked in one or other of the muscle pairs and incorporated into the locomotor pattern. 3. In the majority of sites, the stimulation evoked excitatory responses in the flexor muscles of each of the four limbs during that period of the step cycle in which each respective muscle was naturally active; stimulation in the stance phase of locomotion, although less effective, was also capable of producing responses in these muscles. All three ipsilateral extensor muscles studied [long and lateral heads of triceps and vastus lateralis (Tri, TriL, and VL, respectively)] were normally inhibited during their phase of muscle activity, although excitatory responses were occasionally seen. Responses in the contralateral (co) Tri were invariably excitatory and were largest during the period of muscle activity, whereas responses during the period of activity of the coVL were mixed, with both excitatory and inhibitory responses being seen from any one locus. 4. Excitatory responses were normally largest when stimulation was applied during the time that the muscle was active during the locomotor cycle. Responses evoked at times when the muscle was inactive were sometimes larger than those evoked with the animal at rest; such responses were most commonly seen in the hindlimb flexors and in the coVL. 5. In both flexors and extensors of each of the four limbs, the latency of the responses was greatest when the cat was at rest and least for stimuli given during the period of activity of the respective muscle. Average latencies during the period of muscle activity ranged from a minimum of 9.0 +/- 2.6 (SD) ms for inhibitory responses in the ipsilateral Tri and TriL to a maximum of 17.1 +/- 3.0 ms for the responses evoked in the ipsilateral semitendinosus.(ABSTRACT TRUNCATED AT 400 WORDS)
Swan-Ganz catheter-induced massive hemoptysis and later pulmonary artery false aneurysm occurred in a patient with prosthetic mitral regurgitation. This patient was successfully managed by double-lumen endotracheal intubation, control of pulmonary hypertension, reversal of anticoagulation, mitral valve re-replacement, and transcatheter embolization. The pertinent literature is reviewed.
1. The present article described the various patterns of movement evoked in the limbs and neck by microstimulation (33-ms trains, 330 Hz, 0.2-ms pulses at less than or equal to 35 microA) of the medullary reticular formation (MRF) of seven chronically implanted, unanesthetized, intact cats. Altogether 878 loci were stimulated in 83 penetrations. However, as stimulation in the more lateral regions of the MRF was less effective, the results are based on stimulation in 592 loci made in 56 penetrations at distances of between 0.5 and 2.5 mm lateral to the midline. 2. Of these 592 loci, movement of one or more parts of the body was evoked from a total of 539 (91%) sites. Most of these movements were compound in nature, involving movement of one or more limbs as well as the head. Discrete movements were observed only with respect to the head; limb movements were always accompanied by head movement. In addition, hindlimb movements were always accompanied by forelimb movements, although the inverse was generally not true. 3. The most common effects of the stimulation were as follows: a turning of the head to the ipsilateral side (79% of stimulated sites); flexion of the ipsilateral elbow (41%); and extension of the contralateral elbow (45%). Effects in the hindlimbs were more variable and less frequent, with the majority of the effective loci causing flexion of the ipsilateral knee (9%) together with extension of the contralateral knee (8%). In total, including both flexion and extension, 18% of the stimulated sites caused movement of the ipsilateral hindlimb and 11% of the contralateral hindlimb. 4. Although movements of the head were obtained from the whole extent of the brain stem, movements of the forelimbs showed a dorsoventral organization with flexion of the ipsilateral elbow being evoked from the more dorsal regions of the brain stem, whereas contralateral elbow extension was evoked more frequently from the ventral regions. There was a large area of overlap from which movements of both limbs could be obtained simultaneously. Movements of the hindlimbs were more frequently evoked from central and ventral areas of the brain stem and from the most rostral aspect of the explored region. 5. In examining the combinations of movements evoked by the MRF stimulation, it was found that the most commonly evoked pattern was movement of the head to the stimulated side together with flexion of the ipsilateral forelimb and extension of the contralateral forelimb (26.5% of sites).(ABSTRACT TRUNCATED AT 400 WORDS)
1. The present study has examined the detailed organization of the medullary reticular formation (MRF) as revealed by microstimulation (33-ms trains of 0.2-ms duration pulses at 330 Hz and 35 microA or less) in the intact, chronically implanted, unanesthetized cat. Stimulus-locked electromyographic (EMG) responses were recorded from flexors and extensors of each of the four limbs, as well as bilaterally from muscles of the neck and back, during stimulation of the same 592 loci that formed the basis of the preceding article. 2. The thresholds of the responses were different for each group of muscles, with, on the average, the neck muscles being activated at the lowest range of currents, 13.8-16.5 microA; forelimb muscles at 16.9-17.9 microA; back muscles at 25.4-25.7 microA; and hindlimb muscles at 21.1-25.7 microA. 3. Whereas stimulation within the MRF evoked movement of the head only to the stimulated side (preceding article), analysis of the EMG responses showed there was frequently bilateral activation of the neck muscles. Similarly, even though stimulation produced predominantly ipsilateral elbow flexion and contralateral elbow extension, most loci caused cocontraction of antagonistic muscles at these joints. Cocontraction was also frequently observed for the hindlimbs. Reciprocal activation of antagonistic muscles was less frequent but was observed in the ipsilateral forelimb as well as in both hindlimbs; it was never observed in the contralateral forelimb. 4. Although excitatory responses were observed from widespread regions for all of the muscles under study, those regions of the MRF that evoked the strongest responses in each muscle showed a large degree of segregation. Muscles of the ipsilateral forelimb were most strongly activated from the rostrodorsal MRF, whereas muscles of the contralateral forelimb were most strongly effected by stimulation caudoventrally. Muscles of the hindlimbs were more strongly activated from the rostral brain stem, although with some exceptions. Responses in axial muscles were evoked from widespread regions of the brain stem but were concentrated further caudally than were the limb muscles. 5. Excitatory responses were much more prevalent than inhibitory responses and were evoked from all regions of the MRF, including the most caudal and ventral areas. The shortest latency responses in each track were, on the average, as follows: 6.6-8.8 ms for the neck; 11.2-13.4 ms for the forelimbs; 13.8-14.2 ms for the back; and 15.9-17.2 ms for the hindlimbs. Inhibitory responses were also evoked from widely distributed regions, which were intermingled with those loci evoking excitatory responses.(ABSTRACT TRUNCATED AT 400 WORDS)
Kinematic and electromyographic data were recorded together with motor cortical cell discharge during a task which required the cat to modify its gait in order to step over 3 different types of obstacles fixed to a moving treadmill belt. In order to negotiate the obstacles the cat made large adjustments in limb trajectory which were associated with equally large changes in forelimb flexor muscle activity. Sixteen of 57 identified pyramidal tract neurones recorded from area 4 of two cats increased their peak discharge rate during this gait adjustment. It is suggested that the motor cortex plays a role in adjusting the flexor muscle activity to the requirements of the locomotor task.
To assess the recuperation of locomotor functions of the hindlimbs after spinal section at T13, EMGs and kinematics were analyzed in a chronically implanted adult cat before and up to 14 days after spinalization. At this time, the cat was capable of coordinated walking, plantar foot contact and weight support of the hindquarters for several step cycles. There was however a foot drag early in swing and the steps tended to be shorter. Most muscles showed similar timing characteristics although there were some changes in amplitude. It is concluded that the spinal cord can generate a detailed locomotor pattern in the hindlimbs even in the adult animal.
A kinematic and electromyographic (EMG) analysis was undertaken of the responses evoked in the forelimb of the cat by either mechanical obstruction of the forelimb during the swing phase of locomotion or by electrical stimulation of low-threshold cutaneous afferents during both swing and stance. Mechanical obstruction of the forelimb with a stiff metal rod evoked a complex response that allowed the cat to smoothly negotiate the obstacle without undue disruption of the overall locomotor rhythm. The initial movements were a flexion of the shoulder, together with a locking of the elbow joint, and a dorsiflexion of the wrist, which caused the limb to withdraw from the obstacle. They were followed by an extension of the shoulder, a flexion of the elbow, and a ventroflexion of the wrist, which together brought the limb forward and above the obstacle. The associated and complex pattern of short- and long-latency EMG responses was shown to be related to different aspects of the movement. At the shoulder there was a strong activation of flexor muscles; these responses were of long duration (greater than or equal to 100 ms) and generally lasted throughout the period of shoulder flexion. At the elbow, both flexor and extensor muscles were activated at short latency (9-13 ms). In flexors, this was followed by a cessation and subsequently an augmentation and prolongation of their activity. Dorsiflexors of both the wrist and digits were activated at short latency (10-12 ms) and remained active throughout the period of dorsiflexion of these joints. An injection of a local anesthetic into the area of skin contacted by the metal rod reduced or abolished all of the reflex responses, which suggests that the integrity of cutaneous reflex pathways is essential for the elaboration of these responses. Electrical stimulation of a cutaneous nerve innervating the distal forelimb (the superficial radial nerve) resulted in qualitatively similar, although weaker, responses to those obtained with the mechanical stimulation. Terminal experiments confirmed that these responses were mediated by low-threshold cutaneous afferents. Electrical stimulation also evoked short-latency excitatory responses (10-12 ms) in extensor muscles of the elbow. Generally, the largest reflex effects were obtained during the period of swing for flexor, extensor, and bifunctional muscles. During stance the stimulus was normally ineffective in exciting flexor muscles and in extensors evoked a short-latency inhibition, which was frequently followed by an increase in activity.(ABSTRACT TRUNCATED AT 400 WORDS)
Recordings were made from single units in the medullary reticular formation (MRF) between AP-4.2 and AP-12.9 and from the midline to 3.7 mm lateral in chronically prepared, unrestrained cats walking on a treadmill. Recordings were made with rigid microelectrodes held in a microdrive, and reticulospinal neurons were identified by antidromic stimulation of their axons through microwires chronically implanted into the spinal cord at the L2 level. Electromyograms (EMGs) were recorded from flexor and extensor muscles of the fore- and hindlimbs as well as from back and neck muscles. In total, 295 cells were recorded from 40 penetrations in 4 cats; 252 of these cells were recorded from the more medial regions of the reticular formation encompassing the gigantocellular, magnocellular, and lateral tegmental fields; 38.5% of these (97/252) were antidromically identified from the spinal cord. The remaining 43 neurons (43/295) were recorded from a more lateral and ventral position. These medial and ventrolateral groups of neurons differed not only in position but also in aspects of their discharge during locomotion. Rank-ordered raster displays, triggered from the onset of each recorded muscle, were used to correlate neuronal and muscular activity. The discharge rate of 31% of the reticulospinal neurons (30/97) was modulated once or twice in each step cycle and was strictly related to one or more of the recorded EMGs (EMG-related neurons) on the basis of the pattern of discharge. The discharge of 33/97 (34%) of the neurons was modulated at the periodicity of the locomotor rhythm but could not be correlated with any of the recorded EMGs (locomotor-related cells), whereas the remaining 34/97 neurons (35%) were either silent, fired tonically, or were not related to the locomotor pattern (unrelated cells). Of the EMG-related neurons 27% were related to flexor muscles and the remaining 63% to extensor muscle activity. The discharge pattern of all except two of the flexor-related neurons was correlated with hindlimb muscle activity, whereas that of the extensor-related neurons was correlated almost equally with fore- and hindlimb muscles. Correlations were found with muscles lying both ipsilaterally and contralaterally to the site of the recordings. Although the locomotor-related neurons showed no preferential relation with any of the recorded EMGs, a comparison of the depth of modulation of their discharge measured from postevent histograms suggested that more of these cells were related to the forelimb than to the hindlimb.(ABSTRACT TRUNCATED AT 400 WORDS)
The reflex responses of forelimb muscles to electrical stimulation of the cutaneous superficial radial nerve were recorded during treadmill locomotion in chronically implanted cats. In brachialis and cleidobrachialis (muscles which are active mainly during the swing phase) the responses were maximal during the swing phase and minimal or absent during the stance phase. In the long head of triceps which is active mainly during the stance phase, responses were also minimal during stance and maximal during swing. It is concluded that, for some muscles, the period of maximal reflex responsiveness can be out of phase with the period of the step cycle during which they are normally active.
Chronically implanted microwires were used to deliver brief trains of electrical stimuli (11 cathodal pulses at 330 Hz and intensity 5-35 microA) to sixty-two locations in the grey matter of the pericruciate cortex in cats. Electromyographic (e.m.g.) responses in the contralateral forelimb were recorded from a total of ten muscles (four to eight in each animal) acting about the shoulder, elbow and wrist and on the digits. The animals were relaxed with little background e.m.g. in the muscles and as a result only excitatory effects could be described. Five muscles which are flexors in the locomotor context were excited from more electrodes, distributed more widely across the motor cortex, than another five muscles which are extensors during locomotion; this difference in 'accessibility' was present both at 35 microA stimulus intensity and at 15 microA. At a stimulus intensity of 15 microA, effective cortical electrodes tended to cluster either in the most lateral part of the anterior sigmoid gyrus (rostromedial focus) or in the coronal gyrus just caudal to a line prolonged beyond the lateral end of the cruciate sulcus (caudolateral focus). This is consistent with the existence of a double motor representation within the forelimb motor cortex (Pappas & Strick, 1981). The two foci were similar in that both gave rise to more flexor than extensor responses and to fewer responses in digit or wrist muscles than in muscles acting about more proximal joints (elbow and shoulder). At stimulus intensity 35 microA the latency of the earliest e.m.g. responses ranged from 11 to 14 ms in different muscles. For some muscles and electrodes the amplitude of the e.m.g. responses was substantially altered by a quite small postural change. After pyramidectomy the cortical thresholds and the e.m.g. latencies were both greatly increased.
The forelimb motor cortex was stimulated via chronically implanted microelectrodes whilst electromyographic (e.m.g.) responses were recorded from muscles in the contralateral forelimb in cats walking steadily at 0.5 m/s. The stimuli were brief trains of 0.2 ms pulses (11 pulses at 330 Hz), intensity 5-20 microA and e.m.g.s were recorded from the following muscles: biceps brachii, brachialis, long and lateral heads of triceps brachii, latissimus dorsi, cleidobrachialis, extensor digitorum communis, palmaris longus and flexor and extensor carpi ulnaris. During locomotion, stimulation at 20 microA readily elicited brief, short-latency changes in the normal locomotor patterns of activity in all muscles studied. The changes included production of e.m.g. at times in the step cycle when the muscles are normally inactive and brief augmentations or diminution of the normal locomotor e.m.g.s. Individual electrodes usually influenced several muscles, and muscles acting antagonistically about the same joint were sometimes co-contracted. The first effect on locomotor flexor muscles (i.e. muscles active in relation to the swing phase of the step cycle) was almost always excitatory and such effects were often phase-dependent, usually occurring when the muscle was normally active or about to become active. Extensor muscles were excited from some cortical loci but inhibited from others (inhibitions were necessarily detectable only when the muscles exhibited locomotor-related e.m.g.s). Some micro-electrodes elicited excitation during swing (when the extensors are inactive) but elicited inhibition during stance. In several muscles the latencies of the excitatory e.m.g. changes could be as short as 6 ms measured from the first pulse in the stimulus train. In flexors, but not in extensors, latencies fluctuated according to the timing of the stimuli relative to the step cycle. Reduction in stimulus intensity reduced the amplitude of the e.m.g. changes, the number of muscles influenced and often increased the latency. However, both excitations and inhibitions were sometimes evident at 5 microA and thresholds for excitatory responses were, over-all, substantially lower than in the resting animal. Longer trains of stimuli were capable of resetting the step cycle. Response thresholds were greatly increased after pyramidectomy. These findings support the view that the natural bursts of impulses discharged by pyramidal tract neurones during steady locomotion are likely to contribute to regulating forelimb muscle activity on a step-by-step basis.
Explore the source record for details and available documents.
A method is described for chronically implanting fine flexible microwires into cat motor cortex, which permitted extracellular recordings to be made from 165 single neurones. Most units were recordable for 12 h and some for up to 2 days. Of the neurones tested, 57% were shown to project to the medullary pyramid (pyramidal tract neurones, p.t.n.s). Antidromic latencies corresponded to a range of conduction velocities from 63 to 9 m/s. In the animal at rest neurones discharged at rates from 0.5 to 44 impulses/s. During locomotion at 0.5 m/s (a slow walk) 56% of cells discharged faster than at rest and 80% showed frequency modulations time-locked to the step cycle. Most fired one discrete burst of impulses per step or one peak period superimposed on a maintained discharge. In different cells peak activity occurred at widely different times during the step cycle. A few cells peaked twice per step. Peak rates (averaged over twenty steps) ranged from 10 to over 120 impulses/s, the values for most slow-axon p.t.n.s (conduction velocity less than 21 m/s) being lower than for any of the fast-axon p.t.n.s. For locomotion at speeds between 0.37 and 1.43 m/s a roughly linear relationship existed between discharge rate and speed in 14% of cells. However, the changes were modest and in most cells both mean rate and peak rate were unrelated to speed. In some cells discharge phasing was fixed (relative to the step cycle in the contralateral forelimb); in others there were progressive phase shifts (or more complex changes) as speed increased. During locomotion up a 10 degrees incline discharge phasings were the same as on the flat in all of the twenty-seven neurones studied and most showed no substantial change in mean rate or peak rate (although there were substantial increases in limb muscle electromyogram amplitudes).
Discharge patterns of motor cortical neurones in cats walking steadily on a moving belt have been compared with other functional characteristics of the neurones. In forelimb motor cortex rhythmic discharges occurred in cells with peripheral receptive fields in all parts of the contralateral forelimb and also in cells with no discernible receptive field. Cells discharging at similar times during the step cycle often had very different receptive fields and cells with similar receptive fields (including neighbouring cells) could discharge at similar or at quite different times. In cells with a cutaneous receptive field including the forefoot the discharges during locomotion remained rhythmic (and their phasing relative to the step cycle was unchanged) when the response to mechanical stimulation in the receptive field was temporarily much reduced or abolished by local anaesthesia of the skin. The proportion of neurones showing accelerated firing during different parts of the step cycle fluctuated more for antidromically identified pyramidal tract neurones (p.t.n.s) than for non-p.t.n.s and was highest during the second half of stance in the contralateral forelimb and lowest during swing. When the neurones were subdivided according to the movement evoked by threshold electrical stimulation through the micro-electrode, p.t.n.s and non-p.t.n.s recorded by electrodes evoking elbow flexion showed a wide variety of discharge patterns. For p.t.n.s the discharge rate reached an average of 69 impulses/s during late stance and declined to an average of 26 impulses/s during swing.
Microwires chronically implanted in the pericruciate cortex of free-to-move cats were used to record extracellularly from cortical neurones and to deliver intracortical stimulation. Natural stimulation of cutaneous and/or deep mechanoreceptors in limbs and trunk evoked discharges in 89% of 165 neurones, 57% of which were pyramidal tract neurones. Out of 112 cells with receptive fields on the contralateral forelimb, 41% had cutaneous fields, 29% had fields involving deep tissues and 30% were driven from both sources. Cutaneous receptive fields were much commoner than deep ones among cells with fields including the forefoot; this relationship was reversed for cells with more proximal fields. Many more cells had distal than proximal fields. The 'zones' of the forelimb (i.e. foot, wrist, elbow, shoulder) provided input to widespread and overlapping cell populations within the coronal gyrus and the lateral parts of the anterior and posterior sigmoid gyri. Despite the overlap a somatotopy existed with successively more distal limb zones represented successively further laterally in the pericruciate area. Intracortical stimulation (eleven cathodal pulses, duration 0.2 ms, frequency 330 Hz, intensity 35 microA or less) evoked flick movements of the contralateral limbs which were abolished by pyramidectomy. In the forelimb, shoulder movements were commonest and elbow, wrist and digits were represented with decreasing frequency. Both for 35 microA and for threshold stimulation the distributions of the effective electrodes revealed an overlapping somatotopy such that the wrist movements were almost restricted to the coronal gyrus and shoulder movements were most often evoked from the lateral part of the anterior sigmoid gyrus. The movement and receptive field somatotopies overlapped heavily but the former showed a distinct lateral shift relative to the latter. As a result shoulder movements were not uncommonly evoked from the coronal gyrus although the shoulder provided almost no input to cells in that area.
Electromyographic and kinematic responses of all four limbs were studied when loci within the medullary reticular formation (MRF) were stimulated (30-ms train of 0.2-ms pulses at 300 Hz, strength 35 microA) during treadmill locomotion in spontaneously walking thalamic cats. Responses could be evoked in flexor or extensor muscles of any given limb by such stimulation, depending on the time during the step cycle at which the stimulus was delivered. Stimulation normally excited flexor muscles but could either excite or inhibit extensor muscles depending on the exact position of the electrode. Excitatory responses in extensor muscles were often followed by a short period of inhibition of activity. The responses in muscles of the opposing limbs of the same girdle were, in general, reciprocally organized. For instance, a stimulus delivered during the swing phase of the ipsilateral limb normally evoked excitatory responses both in flexor muscles of that limb and in extensor muscles of the contralateral limb. The same stimulus delivered during the stance phase of the ipsilateral limb evoked excitatory responses in ipsilateral extensor muscles and in contralateral flexor muscles. Responses were also observed at the same time in fore- and hindlimbs that were well organized with respect to the locomotor cycle. Seventy-five percent of all responses occurred within 8-20 ms of the onset of the stimulus train. Responses evoked in muscles of the opposing limbs of one girdle (e.g., a flexor of one limb and an extensor of the other) had similar latencies, suggesting that the responses were synchronously organized on both sides of the body rather than one being a consequence of the other. Although the majority of responses in a given muscle were elicited during its period of activity, responses could occasionally be evoked when there was no activity in that muscle or could be absent despite activity in the muscle. The short trains of stimuli were normally potent enough to affect the limb trajectory, which reflected changes in the onset or the offset of the activity of most muscles. Thus the stimuli effectively changed both the duration of the period of activity in these muscles and the overall step cycle. Longer trains of stimuli (200 ms) markedly amplified these changes to the point of completely resetting the locomotor rhythm.(ABSTRACT TRUNCATED AT 400 WORDS)