Magnetic stimulation of the human motor cortex evokes skin sympathetic nerve activity.

Single-pulse magnetic coil stimulation (Cadwell MES 10) over the cranium induces without pain an electric pulse in the underlying cerebral cortex. Stimulation over the motor cortex can elicit a muscle twitch. In 10 subjects, we tested whether motor cortical stimulation could also elicit skin sympathetic nerve activity (SSNA; n = 8) and muscle sympathetic nerve activity (MSNA; n = 5) in the peroneal nerve. Focal motor cortical stimulation predictably elicited bursts of SSNA but not MSNA; with successive stimuli, the SSNA responses did not readily extinguish (94% of discharges to the motor cortex evoked SSNA responses) and had predictable latencies [739 +/- 33 (SE) to 895 +/- 13 ms]. The SSNA responses were similar after stimulation of dominant and nondominant sides. Focal stimulation posterior to the motor cortex elicited extinguishable SSNA responses. In three of six subjects, anterior cortical stimulation evoked SSNA responses similar to those seen with motor cortex stimulation but without detectable movement; in the other subjects, anterior stimulation evoked less SSNA discharge than that seen with motor cortex stimulation. Contrasting with motor cortical stimulation, evoked SSNA responses were more readily extinguished with 1) peripheral stimulation that directly elicited forearm muscle activation accompanied by electromyograms similar to those with motor cortical stimulation; 2) auditory stimulation by the click of the energized coil when off the head; and 3) in preliminary experiments, finger afferent stimulation sufficient to cause tingling. Our findings are consistent with the hypothesis that motor cortex stimulation can cause activation of both alpha-motoneurons and SSNA.     

Somatosensory evoked potentials following lesions of the claustrum

Ipsi- and contralateral cortical somatosensory evoked potentials (SEP) were recorded following median nerve stimulation in 12 patients with unilateral brain lesions and in 5 healthy subjects. Computed tomographic scans of brain were performed on admission. In all patients with lesions of the claustrum there was absence of SEP contralateral to the side of the lesion and ipsilateral to the stimulated nerve. This phenomenon did not appear in our material following lesions involving other structures e.g. thalamus or somatosensory cortex. Our observations suggest that the claustrum may influence deeply the contralateral somatosensory cortex. This may be due to the fact that a large part of the claustrum is involved in transmission of the sensory information from receptors to the somatosensory cortex.     

Reverberation of recent visual experience in spontaneous cortical waves

Spontaneous waves of activity propagating across large cortical areas may play important roles in sensory processing and circuit refinement. However, whether these waves are in turn shaped by sensory experience remains unclear. Here we report that visually evoked cortical activity reverberates in subsequent spontaneous waves. Voltage-sensitive dye imaging in rat visual cortex shows that following repetitive presentation of a given visual stimulus, spatiotemporal activity patterns resembling the evoked response appear more frequently in the spontaneous waves. This effect is specific to the response pattern evoked by the repeated stimulus, and it persists for several minutes without further visual stimulation. Such wave-mediated reverberation could contribute to short-term memory and help to consolidate the transient effects of recent sensory experience into long-lasting cortical modifications.     

Afferent inhibition and functional properties of neurons in the vibrissae projection area of the cat somatosensory cortex]

The aim of this study was to investigate the role of inhibitory processes in S-1 cortex of cats. The inhibition was evoked by "natural" afferent stimulation of the fascial vibrissae. For this purpose, two neighboring vibrissae were sequentially stimulated by mechanical deflection; single unit activity was recorded simultaneously from the cortex. Results showed that conditioning by afferent stimulation significantly influenced the directional sensitivity of cortical neurons. These data and analysis of spatial pattern of stimulated vibrissa indicate that detector neurons could be quickly modified during sensory processing.     

Spatial attention changes excitability of human visual cortex to direct stimulation

Conscious perception depends not only on sensory input, but also on attention [1, 2]. Recent studies in monkeys [3-6] and humans [7-12] suggest that influences of spatial attention on visual awareness may reflect top-down influences on excitability of visual cortex. Here we tested this specifically, by providing direct input into human visual cortex via cortical transcranial magnetic stimulation (TMS) to produce illusory visual percepts, called phosphenes. We found that a lower TMS intensity was needed to elicit a conscious phosphene when its apparent spatial location was attended, rather than unattended. Our results indicate that spatial attention can enhance visual-cortex excitability, and visual awareness, even when sensory signals from the eye via the thalamic pathway are bypassed.     

Interhemispheric interactions between the human primary somatosensory cortices

In the somatosensory domain it is still unclear at which processing stage information reaches the opposite hemispheres. Due to dense transcallosal connections, the secondary somatosensory cortex (S2) has been proposed to be the key candidate for interhemispheric information transfer. However, recent animal studies showed that the primary somatosensory cortex (S1) might as well account for interhemispheric information transfer. Using paired median nerve somatosensory evoked potential recordings in humans we tested the hypothesis that interhemispheric inhibitory interactions in the somatosensory system occur already in an early cortical processing stage such as S1. Conditioning right S1 by electrical median nerve (MN) stimulation of the left MN (CS) resulted in a significant reduction of the N20 response in the target (left) S1 relative to a test stimulus (TS) to the right MN alone when the interstimulus interval between CS and TS was between 20 and 25 ms. No such changes were observed for later cortical components such as the N20/P25, N30, P40 and N60 amplitude. Additionally, the subcortically generated P14 response in left S1 was also not affected. These results document the existence of interhemispheric inhibitory interactions between S1 in human subjects in the critical time interval of 20-25 ms after median nerve stimulation.     

Respiratory-related cortical potentials evoked by inspiratory occlusion in humans

It has long been recognized that humans can perceive respiratory loads. There have been several studies on the detection and psychophysical quantification of mechanical load perception. This investigation was designed to record cortical sensory neurogenic activity related to inspiratory mechanical loading in humans. Inspiration was periodically occluded in human subjects while the electroencephalographic (EEG) activity in the somatosensory region of the cerebral cortex was recorded. The onset of inspiratory mouth pressure (Pm) was used to initiate signal averaging of the EEG signals. Cortical evoked potentials elicited by inspiratory occlusions were observed when C3 and C alpha were referenced to CZ. This evoked potential was not observed with the control (unoccluded) breaths. There was considerable subject variability in the peak latencies that was related to the differences in the inspiratory drive, as measured by occlusion pressure (P0.1). The results of this study demonstrate that neurogenic activity can be recorded in the somatosensory region of the cortex that is related to inspiratory occlusions. The peak latencies are longer than analogous somatosensory evoked potentials elicited by stimulation of the hand and foot. It is hypothesized that a portion of this latency difference is related to the time required for the subject to generate sufficient inspiratory force to activate the afferents mediating the cortical response.     

Short-term depression at thalamocortical synapses contributes to rapid adaptation of cortical sensory responses in vivo

In vivo whole-cell recordings revealed that during repeated stimulation, synaptic responses to deflection of facial whiskers rapidly adapt. Extracellular recordings in the somatosensory thalamus revealed that part of the adaptation occurs subcortically, but because cortical adaptation is stronger and recovers more slowly, cortical mechanisms must also contribute. Trains of sensory stimuli that produce profound sensory adaptation did not alter intrinsic membrane properties, including resting membrane potential, input resistance, and current-evoked firing. Synaptic input evoked via intracortical stimulation was also unchanged; however, synaptic input from the somatosensory thalamus was depressed by sensory stimulation, and this depression recovered with a time course matching that of the recovery of sensory responsiveness. These data strongly suggest that synaptic depression of thalamic input to the cortex contributes to the dynamic regulation of neuronal sensitivity during rapid changes in sensory input.     

Mechanisms of reflex bladder activation by pudendal afferents

Activation of pudendal afferents can evoke bladder contraction or relaxation dependent on the frequency of stimulation, but the mechanisms of reflex bladder excitation evoked by pudendal afferent stimulation are unknown. The objective of this study was to determine the contributions of sympathetic and parasympathetic mechanisms to bladder contractions evoked by stimulation of the dorsal nerve of the penis (DNP) in α-chloralose anesthetized adult male cats. Bladder contractions were evoked by DNP stimulation only above a bladder volume threshold equal to 73 ± 12% of the distension-evoked reflex contraction volume threshold. Bilateral hypogastric nerve transection (to eliminate sympathetic innervation of the bladder) or administration of propranolol (a β-adrenergic antagonist) decreased the stimulation-evoked and distension-evoked volume thresholds by -25% to -39%. Neither hypogastric nerve transection nor propranolol affected contraction magnitude, and robust bladder contractions were still evoked by stimulation at volume thresholds below the distension-evoked volume threshold. As well, inhibition of distention-evoked reflex bladder contractions by 10 Hz stimulation of the DNP was preserved following bilateral hypogastric nerve transection. Administration of phentolamine (an α-adrenergic antagonist) increased stimulation-evoked and distension-evoked volume thresholds by 18%, but again, robust contractions were still evoked by stimulation at volumes below the distension-evoked threshold. These results indicate that sympathetic mechanisms contribute to establishing the volume dependence of reflex contractions but are not critical to the excitatory pudendal to bladder reflex. A strong correlation between the magnitude of stimulation-evoked bladder contractions and bladder volume supports that convergence of pelvic afferents and pudendal afferents is responsible for bladder excitation evoked by pudendal afferents. Further, abolition of stimulation-evoked bladder contractions following administration of hexamethonium bromide confirmed that contractions were generated by pelvic efferent activation via the pelvic ganglion. These findings indicate that pudendal afferent stimulation evokes bladder contractions through convergence with pelvic afferents to increase pelvic efferent activity.     

Somatosensory evoked potentials (SEPs) elicited by magnetic nerve stimulation

Magnetic stimulation of peripheral nerves at distal and proximal sites of the upper and lower extremities and at the midlumbar level were used to elicit cortical somatosensory evoked potentials. Evidence is provided that peripheral nerve trunks, rather than distal receptor afferents, are the anatomical structures stimulated by the electromagnetic fields. Magnetic stimulation of peripheral nerves is considered to be useful for an evaluation of the integrity of proximal nerves, nerve roots and central conduction along sensory pathways. In contrast to electrical nerve stimulation, magnetic stimulation is painless and can be applied to proximal nerves and plexus. By means of proximal nerve stimulation central sensory conduction can be tested even in patients with peripheral nerve lesions or polyneuropathy.     

Reticular activation by splanchnic afferents: its influence on visual projections at the level of the cerebral cortex in cats]

In an previous work, we have shown a strong increase of visual evoked potentials of the medial lateral gyrus by splanchnic stimulation. This present study demonstrates that this effect is due to the activation of the substantia reticularis. Actually, we observed: 1) an increase of the reticular visual potential by splanchnic stimulus in convergent zone; 2) an increase of the visual evoked potential in the primary cortical area by stimulation of this reticular formation. These results show the existence of a functional pathway including the splanchnic nerve, the reticular formation and the visual cortex.     

Analysis of the organization of afferent inputs in the projection cortex

Fast fluctuations in the evoked potentials (EP) at a local point of the projection cortex following stimulation of different pathways may reflect the activity of pyramidal neurons of different cortical layers. Analysis shows that the afferent and interarea projections to the somatic sensory cortex terminate on different neurons which can be regarded as relay neurons for a given pathway. Each group of neurons has its own system of inhibition for selective control of impulses coming along this pathway at the cortical level.Institute of Normal and Pathological Physiology, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 2, No. 4, pp. 368–372, July–August, 1970.     

Modality-specific organization for cutaneous and proprioceptive sense in human primary sensory cortex studied by chronic epicortical recording

Modality specificity of human primary somatosensory cortex was studied by recording somatosensory evoked potentials (SEPs) from subdural electrodes in a patient with intractable focal motor seizure. A newly developed device was used for selectively activating proprioception. The spatial and temporal distributions of proprioception-related SEPs elicited by brisk passive flexion movement at the proximal interphalangeal (PIP) joint of the middle finger (4 degrees in 25 ms) were quite different from those to cutaneous sense evoked by electric stimulation of the digital nerve at the same site. It was for the first time demonstrated that proprioception-related SEPs following passive finger movement do not originate in area 3b, which was clearly activated by cutaneous stimulation, and that other sites at the sensorimotor cortex such as areas 2, 3a and 4 possibly contribute to the cortical processing of proprioception.     

Effect of injury to the lateral reticular nucleus and nuclei of the inferior olive on electrical responses of the cerebellar cortex evoked by vagus nerve stimulation in cats

The role of the lateral reticular nucleus and nuclei of the inferior olive in the formation of cerebellar cortical evoked potentials in response to vagus nerve stimulation was determined in experiments on 28 cats anesthetized with chloralose and pentobarbital. After electrolytic destruction of the lateral reticular nucleus, in response to vagus nerve stimulation, especially ipsilateral, lengthening of the latent period and a decrease in amplitude of evoked potentials were observed; after bilateral destruction of this nucleus, evoked potentials could be completely suppressed. It is concluded that the lateral reticular nucleus relays interoceptive impulses in the vagus nerve system on to the cerebellar cortex. Additional evidence was given by the appearance of spike responses of Purkinje cells, in the form of mainly simple discharges, to stimulation of the vagus nerve. After destruction of the nuclei of the inferior olive, the latent period and the number of components of evoked potentials in response to vagus nerve stimulation remained unchanged but their amplitude was reduced. The role of the nuclei of the inferior olive as a regulator of the intensity of the flow of interoceptive impulses to the cerebellum is discussed.N. I. Pirogov Medical Institute, Vinnitsa. Translated from Neirofiziologiya, Vol. 9, No. 3, pp. 290–299, May–June, 1977.     

Possible mechanisms of the anterior limbic cortex influence on the neuron activity of the vago-solitary complex

In ananesthetized cats, neurons of the nucleus of the tractus solitarius (NTS) and the dorsal motor nucleus of the vagus nerve (DMNV) revealed phasic excitatory responses to separate single vagal and cortical stimuli. Stimulation of the anterior limbic cortex combined with vagal stimulation resulted in inhibitory or excitatory modification of the vagal induced responses of the NTS and DMNV neurons. The data obtained suggest that complete inhibitory effects are related to general cortical mechanisms of control of the functional state of the brain stem visceral neurons. Selective inhibition of the vagal induced responses by limbic cortex stimulation is due to particular cortical mechanisms of the visceral sensory transmission control via the NTS neurons.     

Amygdaloid or cortical facilitation of antidromic activity of trigeminal motoneurons in the rat

1. Electrical stimulation of the rat's contralateral central amygdaloid (CAm) nucleus or the contralateral frontal cortex markedly augmented the antidromic field potential evoked by stimulation of mylohyoid (Myl) nerve. 2. This facilitation was shown to be due to EPSPs of the mylohyoid-anterior digastric (Myl-Dig) motoneurons. 3. In a few motoneurons, cortical EPSPs had fixed short latencies following high-frequency double stimuli and this is believed to be due to a monosynaptic pathway. 4. The amygdaloid or cortically evoked EPSPs relieved IS-SD blockade in a few motoneurons and also facilitated antidromic discharge in others which did not show any IS or M spike response to the same subthreshold antidromic stimulation. The underlying mechanisms are discussed.     

Somatotopy of human hand somatosensory cortex as studied in scalp EEG

We recorded somatosensory evoked potentials (SEPs) in scalp EEGs during stimulation of the median nerve, the ulnar nerve and the individual digits in 3 normal subjects and in 1 epilepsy patients. In this patient we also measured SEPs from chronically indwelling subdural grid electrodes during electrocorticography (ECoG). We applied dipole modelling techniques to study the 3-dimensional intracerebral locations and time activities of the neuronal sources underlying stimulation of different peripheral receptive fields. The sources underlying median nerve SEPs were located an average of 10.8 mm lateral inferior to those underlying ulnar nerve SEPs. Digit SEP sources showed a somatotopic arrangement from lateral inferior to medial superior in the order thumb, index finger, middle finger, ring finger and little finger, with some overlap or reversal for adjacent digits. The average distance between thumb and little finger was 12.5 mm. Thumb, index finger and middle finger were clustered around median nerve cortical representation, whereas ring finger and little finger were arranged around ulnar nerve cortex. In the epilepsy patient, the source localizations obtained in scalp EEGs showed good agreement with those on ECoGs. We conclude that SEPs recorded in scalp EEGs can be used to study functional topography of human somatosensory cortex non-invasively.     

Neuronal mechanisms of sensory integration in the visual analyzer system

It has been established in experiments on immobilized cats that somatic and interoceptive signals produce complex reorganizations of the spontaneous and evoked activity of visual cortex units. Either long diffuse changes of spike frequency or phasic reactions have been observed. The dynamics of sensory integration in the visual cortex is determined by unconditioned and conditioned mechanisms both of an intra- and interanalyser nature. Functioning of the microsystem of learning elements in the visual cortex is based on units capable of fixing the elaborated changes of evoked activity and constituting 18.6 percent of the total number of cellular elements in the visual projection cortex, responding to direct cortical stimulation. Microinophoresis of synaptically active agents has shown that complexely organized choline- and serotoninergic structures involved in the processes of unconditioned and conditioned interaction of heteromodal excitations are located in the visual cortex.     

Activation and inhibition of the micturition reflex by penile afferents in the cat

Coordination of the urinary bladder and the external urethral sphincter is controlled by descending projections from the pons and is also subject to modulation by segmental afferents. We quantified the effects on the micturition reflex of sensory inputs from genital afferents traveling in the penile component of the somatic pudendal nerve by electrical stimulation of the dorsal nerve of the penis (DNP) in alpha-chloralose anesthetized male cats. Depending on the frequency of stimulation (range, 1-40 Hz), activation of penile afferents either inhibited contractions of the bladder and promoted urine storage or activated the bladder and produced micturition. Stimulation of the DNP at 5-10 Hz inhibited distension-evoked contractions and increased the maximum bladder capacity before incontinence. Conversely, stimulation at 33 and 40 Hz augmented distension-evoked contractions. When the bladder was filled above a threshold volume (70% of the volume necessary for distension-evoked contractions), stimulation at 20-40 Hz activated de novo the micturition reflex and elicited detrusor contractions that increased voiding efficiency compared with distension-evoked voiding. Electrical stimulation of the DNP with a cuff electrode or percutaneous wire electrode produced similar results. The ability to evoke detrusor contractions by activation of the DNP was preserved following acute spinal cord transection. These results demonstrate a clear role of genital afferents in modulating the micturition reflex and suggest the DNP as a potential target for functional restoration of bladder control using electrical stimulation.