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Biomedical subjects
D M Armstrong
Publications and source records attributed to D M Armstrong.
Electromyographic responses evoked in muscles of the forelimb by intracortical stimulation in the cat.
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.
Action Potentials↗
Forelimb electromyographic responses to motor cortex stimulation during locomotion in the cat.
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.
Action Potentials↗
Enkephalin-like immunoreactivity in rat area postrema: ultrastructural localization and coexistence with serotonin.
The ultrastructure of enkephalin-containing neurons and their capacity to take-up [3H]serotonin were examined in the area postrema. Untreated adult rats and rats with intraventricular infusions of 10(-4) M tritiated serotonin, 5-hydroxytryptamine [( 3H]5-HT) were perfused with 4% paraformaldehyde and 0.2-0.5% glutaraldehyde. Coronal Vibratome sections through the area postrema from both groups of animals were immunocytochemically labeled with an antiserum to leucine Leu5-enkephalin. The sections from the animals infused with the isotope subsequently were processed for autoradiography. Enkephalin-like immunoreactivity (ELI) was detected in perikarya, dendrites, axons and axon terminals most frequently located along the ventricular and ventrolateral portions of the area postrema. The labeled perikarya were few in number and were characterized by a thin rim of cytoplasm containing peroxidase immunoreactivity. Dendrites and terminals containing ELI formed synapses primarily with unlabeled axon terminals and dendrites, respectively. However, terminals containing ELI also formed synaptic junctions with other unlabeled axon terminals. Appositions between enkephalin-containing processes and modified glia were occasionally seen near the ventricular surface. In sections processed for both immunocytochemistry and autoradiography, approximately 5% of the terminals containing ELI showed uptake of [3H]5-HT. We conclude that neurons containing ELI are primarily, but not exclusively, associated with other intrinsic neurons or afferents in the rat area postrema and that some of the enkephalin-labeled terminals have the capacity to take-up serotonin. Specificity of uptake of [3H]5-HT in neurons containing endogenous serotonin and factors which may contribute to the low probability of detecting both peroxidase and autoradiographic markers in single sections are discussed.
Animals↗
Discharges of pyramidal tract and other motor cortical neurones during locomotion in the cat.
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).
Action Potentials↗
Locomotor-related neuronal discharges in cat motor cortex compared with peripheral receptive fields and evoked movements.
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.
Action Potentials↗
Topographical localization in the motor cortex of the cat for somatic afferent responses and evoked movements.
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.
Animals↗
Discharges of nucleus interpositus neurones during locomotion in the cat.
Extracellular recordings were made from ninety-five cerebellar nuclear neurones in the cat. All were studied during periods of steady walking at 0.5 m/s and most were also studied in the resting animal. Most neurones were in nucleus interpositus anterior; forty-four cells were shown by antidromic invasion to project to the mid-brain. Most neurones discharged tonically in the absence of overt movements and the mean rate was 42 impulses/s (S.D. +/- 23). During locomotion the mean rate was 68 impulses/s (S.D. +/- 32). In all but seven neurones the discharge during locomotion was frequency modulated but in different neurones the depth of modulation varied from 5 to 161 impulses/s (mean 52 impulses/s; S.D. +/- 30) and the time of peak discharge relative to the step cycle in the ipsilateral forelimb also varied widely. Despite the individual differences the population as a whole was much more active during forelimb swing than during stance, both in numbers of neurones strongly active and in over-all average discharge rate (74 impulses/s as compared with 55). Most neurones had tactile receptive fields on the ipsilateral forelimb while others received input from head and neck or from both ipsilateral limbs. The tendency to discharge preferentially during early swing was greatest for the first group, especially the subpopulations with receptive fields around or proximal to the elbow. Cells encountered in close sequence during a micro-electrode track had similarly located receptive fields and usually showed similar patterns of discharge during locomotion. These findings are discussed in relation to the suggestion by Orlovsky (1972a, b, c) that nucleus interpositus assists in regulating locomotion by evoking rubrospinal discharges which facilitate the flexor muscle activities produced by the spinal mechanisms responsible for generating the swing phase of the step cycle.
Action Potentials↗
Discharges of Purkinje cells in the paravermal part of the cerebellar anterior lobe during locomotion in the cat.
Extracellular recordings were made from 124 Purkinje cells in the paravermal part of lobule V of the cerebellum in cats walking steadily at a speed of 0.5 m/s on a moving belt. All cells tested had a tactile receptive field from which simple spikes could be evoked and 96% of these were on the ipsilateral forelimb. Seventy-six of the cells were also studied whilst the animals sat or lay quietly without movement. Complex spikes were discharged at 1-2/s and these were accompanied by simple spikes in fifty-nine cells (78%); in the remaining cells there were no or few simple spikes. The over-all mean discharge rate (including both types of spike) was 37.8 +/- 27 impulses/s (+/- S.D.). During locomotion all cells discharged both types of spike and the over-all mean rate was 57.6 +/- 29 impulses/s (+/- S.D.). In all cells but one, the frequency of the simple spikes was modulated rhythmically in time with the stepping movements but the phasing relative to the step cycle varied widely between cells. Peak rates also varied widely, the average being 91.5 +/- 44 impulses/s (+/- S.D.). Most cells (63%) generated one period of accelerated discharge per step but others generated two (35%) or three (2%) such periods. Despite the individual variations in discharge timing the population as a whole was considerably more active during the swing than the stance phase of the step cycle in the ipsilateral forelimb (68 impulses/s as compared with 49 impulses/s on average). Thirty-four cells were electrophysiologically identified as lying in the c1 zone of the cortex and twenty-five as being in the c2 zone (nomenclature of Oscarsson, 1980). During locomotion, the population activity in the two zones differed slightly: activity in the c1 population was phase advanced by approximately one-tenth of the step cycle. The results are discussed, with particular emphasis on the finding that population activity in the Purkinje cells of the c1 zone fluctuated during the step cycle in parallel with that in the part of nucleus interpositus to which they project.
Action Potentials↗
Ultrastructural localization of monoamines and peptides in rat area postrema.
An overall schema for the synaptic interactions of monoaminergic and peptidergic neurons and their relation to the ventricle and to blood vessels within the rat area postrema is presented. The specific markers include: 1) the immunocytochemical localization of the catecholamine-synthesizing enzyme tyrosine hydroxylase, and the neuropeptides enkephalin and substance P; and 2) the radioautographic localization of [3H]serotonin (5-hydroxytryptamine) and 3H-labeled amino acids anterogradely transported from the nodose ganglion.
Amines↗
NADPH-diaphorase: a selective histochemical marker for the cholinergic neurons of the pontine reticular formation.
Choline acetyltransferase immunohistochemistry has identified a large group of cholinergic neurons in the pontine tegmentum. By combined immunohistochemical and enzyme histochemical studies this particular cholinergic cell group was found to contain an enzyme, NADPH-diaphorase, that can be visualized histochemically. Thus NADPH-diaphorase histochemistry provides a simple, reliable method to selectively stain the cholinergic neurons of the brainstem reticular formation. The resolution obtained by this novel histochemical technique is similar to that found with the Golgi stain, and it should therefore be of great value in morphological studies of this cholinergic cell group.
Animals↗
The olivocerebellar projection in the rat: an autoradiographic study.
The extent of the olivocerebellar projection was examined in the rat using autoradiographic techniques. In animals in which injections of [3H]leucine encompassed the whole olive unilaterally (4 cases), the vast majority of olive cells was densely labelled and climbing fibres were heavily labelled throughout the contralateral hemicerebellum, except for some small gaps which were not consistently located between cases. The multiple injections required to cover the oliver inevitably labelled cells in the reticular formation surrounding the olive, and it is possible that these neurones might also provide climbing fibres to the cerebellum. To control for this possibility, the inferior olive was pharmacologically destroyed (4 cases) prior to [3H]leucine injections similar in size and placement to those given to normal animals. Examination of the cerebellar cortex of these pretreated animals revealed no molecular layer labelling despite identification of labelled reticular neurones. It was thus demonstrated that all regions of the cerebellar cortex receive afferents from the inferior olive which terminate as climbing fibres. The distribution of these terminations over the entire cortex permits the conclusion that the inferior olive is the major source of climbing fibres in the rat. The same conclusions are reached using [3H]methionine as the tracer (4 cases).
Afferent Pathways↗
Topographical localization in the olivocerebellar projection in the rat: an autoradiographic study.
The topographical organization of the olivocerebellar projection in the rat was examined by making small localized injections of tritiated leucine into the inferior olive (12 cases). The overall projection pattern is displayed in the form of a topographical map, revealing that the olivocerebellar projection is organized in sagittally orientated strips of cerebellar cortex, each of which receives its terminations from a discrete portion of the olive. The projection pattern is compared to that in the cat. In addition, the laterality of the projection is discussed.
Afferent Pathways↗
Distribution of cholinergic neurons in rat brain: demonstrated by the immunocytochemical localization of choline acetyltransferase.
The neuroanatomical location and cytological features of cholinergic neurons in the rat brain were determined by the immunocytochemical localization of the biosynthetic enzyme, choline acetyltransferase (ChAT). Perikarya labeled with ChAT were detected in four major cell groups: (1) the striatum, (2) the magnocellular basal nucleus, (3) the pontine tegmentum, and (4) the cranial nerve motor nuclei. Labeled neurons in the striatum were observed scattered throughout the neostriatum (caudate, putamen) and associated areas (nucleus accumbens, olfactory tubercle). Larger ChAT-labeled neurons were seen in an extensive cell system which comprises the magnocellular basal nucleus. This more or less continuous set of neuronal clusters consists of labeled neurons in the nucleus of the diagonal band (horizontal and vertical limbs), the magnocellular preoptic nucleus, the substantia innominata, and the globus pallidus. Labeled neurons in the pontine tegmentum were seen as a group of large neurons in the caudal midbrain, dorsolateral to the most caudal part of the substantia nigra, and extended in a caudodorsal direction through the midbrain reticular formation into the area surrounding the superior cerebellar peduncle. The neurons in this latter group constitute the pedunculopontine tegmental nucleus (PPT). An additional cluster of cells was observed medially adjacent to the PPT, in the lateral part of the central gray matter at the rostral end of the fourth ventricle. This group corresponds to the laterodorsal tegmental nucleus. Large ChAT-labeled neurons were also observed in all somatic and visceral motor nerve nuclei. The correspondence of the distribution of ChAT-labeled neurons identified by our methods to earlier immunocytochemical and acetylcholinesterase histochemical studies and to connectional studies of these groups argues for the specificity of the ChAT antibody used.
Animals↗
A comparison of the distribution of central cholinergic neurons as demonstrated by acetylcholinesterase pharmacohistochemistry and choline acetyltransferase immunohistochemistry.
The topographical distribution of cholinergic cell bodies has been studied in the rat brain and spinal cord by choline acetyltransferase (ChAT)-immunohistochemistry and acetylcholinesterase (AChE)-pharmacohistochemistry using diisopropylfluorophosphate (DFP). The ChAT-containing cells and the cells that stained intensely for AChE 4-8 hr after DFP were mapped in detail on an atlas of the forebrain (telencephalon, diencephalon) hindbrain (mesencephalon, rhombencephalon) and cervical cord (C2, C6). Striking similarities were observed between ChAT-positive cells and neuronal soma that stained intensely for AChE both in terms of cytoarchitectural characteristics, and with respect to the distribution of the labelled cells in many areas of the central nervous system (CNS). In the forebrain these areas include the caudatoputamen, nucleus accumbens, medial septum, nucleus of the diagonal band, magnocellular preoptic nucleus and nucleus basalis magnocellularis. In contrast, a marked discrepancy was observed in the hypothalamus and ventral thalamus where there were many neurons that stained intensely for AChE, but where there was an absence of ChAT-positive cells. No cholinergic perikarya were detected in the cerebral cortex, hippocampus, amygdala and dorsal diencephalon by either histochemical procedure. In the hindbrain, all the motoneurons constituting the well-established cranial nerve nuclei (III-VII, IX-XII) contained ChAT and exhibited intense staining for AChE. Further, a close correspondence was observed in the distribution of labeled neurons obtained by the two histochemical procedures in the midbrain and pontine tegmentum, including the laterodorsal tegmental nucleus, some areas in the caudal pontine and bulbar reticular formation, and the central gray of the closed medulla oblongata. On the other hand, AChE-intense cells were found in the nucleus raphe magnus, ventral part of gigantocellular reticular nucleus, and flocculus of the cerebellum, where ChAT-positive cells were rarely observed. According to both techniques, no positive cells were seen in the cerebellar nuclei, the pontine nuclei, or the nucleus reticularis tegmenti pontis. Large ventral horn motoneurons and, occasionally, cells in the intermediomedial zone of the cervical cord displayed ChAT-immunoreactivity and intense AChE staining. On the other hand, AChE-intense cells were detected in the dorsal portion of the lateral funiculus, but immunoreactive cells were not found in any portion of the spinal cord white matter.(ABSTRACT TRUNCATED AT 400 WORDS)
Acetylcholine↗
Monoclonal antibodies to choline acetyltransferase: production, specificity, and immunohistochemistry.
Immunohistochemical localization of choline acetyltransferase (ChAT) in cholinergic neurons has been difficult to achieve because of problems encountered in producing specific antisera. Here we describe the production and characterization of several distinct monoclonal antibodies to ChAT. Each of the monoclonal antibodies exhibits one of three general patterns of cross-species reactions; one pattern shows reactivity limited mainly to bovine ChAT, a second pattern shows reactivity only to ChAT from higher mammals including humans, and the third pattern shows reactivity to ChAT from all mammals tested. The antibodies bound specifically to two closely related bovine proteins of 68,000 and 70,000 daltons using the Western blotting technique. One of the antibodies was used to localize immunohistochemically known cholinergic structures in the rat brain, including motor neurons, basal forebrain neurons, and neostriatal neurons.
Animals↗
Distribution of dopamine-, noradrenaline-, and adrenaline-containing cell bodies in the rat medulla oblongata: demonstrated by the immunocytochemical localization of catecholamine biosynthetic enzymes.
The immunocytochemical localization of the biosynthetic enzymes--tyrosine hydroxylase (TH), dopamine-beta-hydroxylase (DBH), and phenylethanolamine-N-methyltransferase (PNMT)--was used to determine the cytological features and precise neuroanatomical location of catecholaminergic neurons in the medulla oblongata of rat. Perikarya labeled with TH were detected in two bilaterally symmetrical columns located in the ventrolateral and dorsomedial medulla. The distribution and the number of neuronal perikarya containing TH were the same as those containing DBH, except in the dorsal motor nucleus of the vagus at the level of the area postrema where the number of neurons immunocytochemically labeled for TH was considerably greater than those labeled for DBH. The detection of perikarya which show immunoreactivity for TH, used in the biosynthesis of dopamine, noradrenaline, and adrenaline, but not DBH, which converts dopamine to noradrenaline, suggests the existence of dopamine-synthesizing neurons in the medulla. Perikarya labeled with PNMT, used in the biosynthesis of adrenaline, were localized in more restricted regions corresponding to rostral subsets of the dorsal and ventral groups labeled for TH and DBH. Counts of neurons immunocytochemically labeled for TH or PNMT were obtained in order to determine the relative ratio of neurons which contain the enzymes necessary for the synthesis of dopamine, noradrenaline, or adrenaline at various levels of the medulla. At the most caudal levels no PNMT labeled neurons were detected. Further rostral, PNMT-labeled neurons were first detected in the ventrolateral medulla. At the level of the area postrema, the number of PNMT-labeled neurons in the ventrolateral medulla was approximately half of the number of cells showing immunoreactivity for TH. In contrast, few PNMT-labeled cells were detected in the dorsomedial medulla at the level of the area postrema compared to many neurons labeled for TH. At rostral medullary levels, in both the ventrolateral and the dorsomedial regions, the number of neurons labeled for TH and PNMT was essentially the same. Thus most, if not all, of the catecholaminergic neurons in the rostral medulla have PNMT, necessary for the synthesis of adrenaline.
Animals↗
Electron microscopic immunocytochemical localization of substance P in the area postrema of rat.
Electron microscopic immunocytochemistry was employed to examine the ultrastructure of processes containing substance P-like immunoreactivity (SP-LI) in the area postrema of rat. SP-LI was present within axons and axon terminals which formed synapses with dendrites of intrinsic neurons in the rat area postrema. Specialized contacts between labeled processes and perivascular or ependymal glia cells were not observed. These data suggested that substance P may act as a neurotransmitter involved primarily with neuronal (axodendritic) interactions within the area postrema.
Animals↗