Galvanic vestibular stimulation (GVS) is a simple, safe, and specific way to elicit vestibular reflexes. Yet, despite a long history, it has only recently found popularity as a research tool and is rarely used clinically. The obstacle to advancing and exploiting GVS is that we cannot interpret the evoked responses with certainty because we do not understand how the stimulus acts as an input to the system. This paper examines the electrophysiology and anatomy of the vestibular organs and the effects of GVS on human balance control and develops a model that explains the observed balance responses. These responses are large and highly organized over all body segments and adapt to postural and balance requirements. To achieve this, neurons in the vestibular nuclei receive convergent signals from all vestibular receptors and somatosensory and cortical inputs. GVS sway responses are affected by other sources of information about balance but can appear as the sum of otolithic and semicircular canal responses. Electrophysiological studies showing similar activation of primary afferents from the otolith organs and canals and their convergence in the vestibular nuclei support this. On the basis of the morphology of the cristae and the alignment of the semicircular canals in the skull, rotational vectors calculated for every mode of GVS agree with the observed sway. However, vector summation of signals from all utricular afferents does not explain the observed sway. Thus we propose the hypothesis that the otolithic component of the balance response originates from only the pars medialis of the utricular macula.

Probing the human vestibular system with galvanic stimulation

The thalamocortical vestibular system in animals and humans

A recent technique of assessing vestibular function, the vestibular-evoked myogenic potential (VEMP), is an otolith-mediated, short-latency reflex recorded from averaged sternocleidomastoid electromyography in response to intense auditory clicks delivered via headphones. Since their first description 10 years ago, VEMPs are now being used by investigators worldwide, and characteristic changes observed with aging and in a variety of peripheral and central vestibulopathies have been described. Additional methods of evoking VEMPs, which use air- and bone-conducted short-tone bursts, forehead taps, and short-duration transmastoid direct current (DC) stimulation, have been described, and these complement the original technique. Click-evoked VEMPs are attenuated or absent in a proportion of patients with vestibular neuritis, herpes zoster oticus, late Meniere disease, and vestibular schwannomas; their amplitudes are increased and thresholds are pathologically lowered in superior semicircular canal dehiscence presenting with the Tullio phenomenon. VEMPs evoked by clicks and DC are useful when monitoring the efficacy of intratympanic gentamicin therapy used for chemical vestibular ablation. Prolonged p13 and n23 peak latencies and decreased amplitudes have been observed in association with central vestibulopathy. VEMPs evoked by clicks are a robust, reproducible screening test of otolith function. DC stimulation enables differentiation of labyrinthine from retrolabyrinthine lesions; bone-conducted stimuli permit VEMP recording despite conductive hearing loss and deliver a relatively larger vestibular stimulus for a given level of auditory perception.

Characteristics and clinical applications of vestibular-evoked myogenic potentials

liams for their clarifi cation on state and national regulations aimed at reducing the risks of salmonellosis and their advice on conducting this survey. We also thank members and staff of the National Association of State Public Health Veterinarians and the National Resource Center for Health and Safety in Child Care and Early Education for their assistance. a population-based, case-control study.nas and Salmonella Marina infection in children: a refl ection of the increasing incidence of reptile-associated sal-monellosis in the United States. 6. Centers for Disease Control and Prevention. Multistate outbreak of human Salmonella Typhimurium infections associated with aquatic frogs—United States, Wild Boars as Hosts of Human-Pathogenic Anaplasma phagocytophilum Variants To the Editor: Michalik et al. (1) reported a 12% prevalence of Ana-plasma phagocytophilum, the caus-ative agent of human granulocytic anaplasmosis and tick-borne fever of ruminants, in wild boars in Poland. A. phagocytophilum has been reported with low prevalence among wild boar in the Czech Republic, Slovenia (2), and Japan (3). In Spain and Missis-sippi, United States, A. phagocyto-philum in wild boars or feral pigs, respectively , has not been reported (4,5). Furthermore, in Slovenia and Poland, the A. phagocytophilum gene sequences found in samples from wild boars were identical to those found in samples from humans and the tick vector Ixodes ricinus (1). These results suggested , as pointed out by Michalik et al. (1), that wild boar might play a role in the epizootiology of A. phagocyto-philum by serving as a natural reservoir host, at least in some regions. To test this hypothesis, we conducted transcriptomics studies to characterize host response to A. phagocy-tophilum infection in naturally and experimentally infected boars (6,7). The results suggested that boars are susceptible to A. phagocytophilum, but are able to control infection, mainly through activation of innate immune responses and cytoskeleton rearrangement to promote phagocy-tosis and autophagy. Control of A. phagocytophilum infection in boars might result in infection levels below PCR detection or infection clearance, contributing to the low percentage of infection prevalence detected for this species in most regions. The low detection levels suggest that boars have a low or no impact as a reservoir host for A. phagocytophilum. Even if boars remain persistently infected with A. phagocytophilum at low levels by downregulating some adap-tive immune genes and delaying the apoptotic death of neutrophils through activation of the Jak-STAT pathway, among other mechanisms (6), their role as a source of infection for …

In References to References

This review considers whether the vestibular system includes separate populations of sensory axons innervating individual organs and giving rise to distinct central pathways. There is a variability in the discharge properties of afferents supplying each organ. Discharge regularity provides a marker for this diversity since fibers which differ in this way also differ in many other properties. Postspike recovery of excitability determines the discharge regularity of an afferent and its sensitivity to depolarizing inputs. Sensitivity is small in regularly discharging afferents and large in irregularly discharging afferents. The enhanced sensitivity of irregular fibers explains their larger responses to sensory inputs, to efferent activation, and to externally applied galvanic currents, but not their distinctive response dynamics. Morphophysiological studies show that regular and irregular afferents innervate overlapping regions of the vestibular nuclei. Intracellular recordings of EPSPs reveal that some secondary vestibular neurons receive a restricted input from regular or irregular afferents, but that most such neurons receive a mixed input from both kinds of afferents. Anodal currents delivered to the labyrinth can result in a selective and reversible silencing of irregular afferents. Such a functional ablation can provide estimates of the relative contributions of regular and irregular inputs to a central neuron’s discharge. From such estimates it is concluded that secondary neurons need not resemble their afferent inputs in discharge regularity or response dynamics. Several suggestions are made as to the potentially distinctive contributions made by regular and irregular afferents: (1) Reflecting their response dynamics, regular and irregular afferents could compensate for differences in the dynamic loads of various reflexes or of individual reflexes in different parts of their frequency range; (2) The gating of irregular inputs to secondary VOR neurons could modify the operation of reflexes under varying behavioral circumstances; (3) Two-dimensional sensitivity can arise from the convergence onto secondary neurons of otolith inputs differing in their directional properties and response dynamics; (4) Calyx afferents have relatively low gains when compared with irregular dimorphic afferents. This could serve to expand the stimulus range over which the response of calyx afferents remains linear, while at the same time preserving the other features peculiar to irregular afferents. Among those features are phasic response dynamics and large responses to efferent activation; (5) Because of the convergence of several afferents onto each secondary neuron, information transmission to the latter depends on the gain of individual afferents, but not on their discharge regularity.

/pdf/afferent-diversity-and-the-organization-of-central-4l0f3xj0cu.pdf

Afferent diversity and the organization of central vestibular pathways

1. Connections from the utricular (UT) nerve to motoneurons and interneurons in the ipsilateral abducens (AB) nucleus were studied in anesthetized and decerebrated cats. Bipolar electrodes were fixed on the left UT nerve under visual observation. The other branches of the vestibular nerve and the facial nerve were transected in the left inner ear. 2. Stimulation of the UT nerve evoked a small positive-negative (P/N) deflection and a negative (N1) potential in the vestibular nuclei, with mean latencies of 0.56 and 0.84 ms, respectively. In the AB nucleus a small P/N deflection with a mean latency of 0.72 ms was recorded, which was considered as a incoming volley of the UT nerve. 3. Excitatory postsynaptic potentials (EPSPs) were recorded from AB motoneurons with short latencies after UT nerve stimulation. They were classified into two types, M and D. M-type EPSPs, which followed repetitive high-frequency stimuli and were recorded from the majority of AB motoneurons, had latencies ranging from 0.9 to 1.2 ms. Double shocks to the UT nerve evoked EPSPs that had the same latency. It was suggested that the AB motoneurons had monosynaptic connections with the UT nerve. D-type EPSPs, which were recorded from most of the AB motoneurons, had slightly longer latencies ranging from 1.2 to 1.8 ms. They showed temporal facilitation when double shocks were provided to the UT nerve. They did not follow repetitive high-frequency stimuli (< or = 2.5-ms interval). It was suggested that D-type EPSPs were di-synaptically evoked via secondary vestibular neurons. Interneurons in the AB nucleus had the same characteristics as AB motoneurons.(ABSTRACT TRUNCATED AT 250 WORDS)

Monosynaptic and disynaptic connections in the utriculo-ocular reflex arc of the cat

The axonal pathway, conduction velocities, and locations of the cell bodies of utricular nerve-activated vestibulospinal neurons were studied in decerebrated or anesthetized cats using the collision test of orthodromic and antidromic spikes. For orthodromic stimulation, bipolar tungsten electrodes were placed on the utricular nerve and the other vestibular nerve branches were transected. Monopolar tungsten electrodes were positioned on both sides of the upper cervical segments (C2–4), caudal end of the cervical enlargement (C7-T1), and from the lower thoracic to the upper lumbar segments (T12-L3) and were used for antidromic stimulation of the spinal cord. Another monopolar electrode was also placed in the oculomotor nucleus to study whether utricular nerve-activated vestibulospinal neurons have ascending branches to the oculomotor nucleus. Of the 173 vestibular neurons orthodromically activated by the stimulation of the utricular nerve, 46 were second-order vestibulospinal neurons and 5 were third-order neurons. The majority of the utricular nerve-activated vestibulospinal neurons were located in the rostral part of the descending vestibular nucleus and the caudal part of the ventral lateral nucleus. Seventy-three percent of the utricular nerve-activated vestibulospinal neurons descended through the ipsilateral lateral vestibulospinal tract. Approximately 80% of these neurons reached the cervicothoracic junction, but a few reached the upper lumbar spinal cord. Twenty-seven percent of the utricular nerve-activated vestibulospinal neurons descended through the medial vestibulospinal tract or the contralateral vestibulospinal tracts. Those axons terminated mainly in the upper cervical segments. Almost none of the utricular nerve-activated vestibular neurons had ascending branches to the oculomotor nucleus.

Properties of utricular nerve-activated vestibulospinal neurons in cats

Neural connections from the saccular and utricular nerves to the ipsilateral vestibular neurons and the commissural effects were studied by using intracellular recordings of excitatory (E) and inhibitory (I) postsynaptic potentials (PSPs) in vestibular neurons of cats after focal stimulation of the saccular and the utricular maculae. Neural circuits from the maculae to vestibular neurons, termed cross-striolar inhibition, may provide a mechanism for increasing the sensitivity to linear acceleration and tilt of the head. It was examined whether secondary vestibular neurons activated by an ipsilateral otolith organ received a commissural inhibition from a contralateral otolith organ that occupied the same geometric plane. Results suggest that utricular-activated vestibular neurons receiving commissural inhibition may provide a mechanism for increasing the sensitivity to horizontal linear acceleration and tilt of the head. The commissural inhibition of the saccular system was much weaker than that of the utricular system.

Cross‐Striolar and Commissural Inhibition in the Otolith System

We studied the convergence of two afferent pairs of single vestibular neurons by selective stimulation of the horizontal semicircular canal (HC) and saccular (SAC) nerves, and the HC and utricular (UT) nerves in decerebrate cats. All recorded neurons were classified as vestibulospinal (VS), vestibulo-oculospinal (VOS) or vestibulo-ocular (VO), by antidromic stimulation from the oculomotor/trochlear nuclei and the spinal cord: neurons that could not be activated from any test sites were classified as vestibular (V) neurons. Of a total of 125 neurons activated by stimulation of the HC/SAC nerves, 21(17%) received convergent inputs. Twelve of 21 neurons received monosynaptic excitatory inputs from both nerves. About half (9/21, 43%) of the convergent neurons were classified as VS neurons, the majority of which descended through the ipsilateral lateral vestibulospinal tract (i-LVST). The HC/SAC convergent neurons were located in the rostral part of the descending, the medial and the caudal-ventral part of the lateral vestibular nucleus. In 80 neurons studied by stimulation of the HC/UT nerves, both inputs converged in 12 (15%) neurons, more than half of which were VS neurons. Eight of 12 convergent neurons received excitatory inputs followed by inhibition from both the HC and UT nerves. A few convergent neurons (3/12) projected to the oculomotor/trochlear nucleus. Half of the convergent and non-convergent VS neurons descended to the spinal cord through the i-LVST, and the only one VOS convergent neuron via the medial vestibulospinal tract. Most of the convergent neurons were located in the lateral, the rostral part of the descending and medial vestibular nucleus. The percentages of HC/SAC and HC/UT convergence were half those of the posterior semicircular canal (PC), PC/SAC (33%) and PC/UT (33%) convergence, respectively. The convergent neurons receiving the HC and otolith inputs may contribute at least partly to the vestibulocollic reflex.

Convergence of the horizontal semicircular canal and otolith afferents on cat single vestibular neurons.

Convergence between posterior canal (PC) and saccular (SAC) inputs in single vestibular nuclei neurons was investigated in decerebrated cats. Postsynaptic potentials were recorded intracellularly after selective stimulation of the SAC and PC nerves. Stimulation of either the SAC or PC nerve orthodromically activated 143 vestibular nuclei neurons. Of these, 61 (43%) were antidromically activated by stimulation of the C1–C2 junction, 14 (10%) were antidromically activated by stimulation of the oculomotor or trochlear nucleus, and 14 (10%) were antidromically activated by stimulation of both the oculomotor or trochlear nucleus and the spinal cord. Fifty-four (38%) neurons were not activated by stimulation of either or both. We named these neurons vestibulospinal (VS), vestibulo-ocular (VO), vestibulo-oculo-spinal (VOS) and vestibular (V) neurons, respectively. Both PC and SAC inputs converged in 47 vestibular nuclei neurons (26 VS, 2 VO, 6 VOS and 13 V neurons). Of these, 19 received monosynaptic excitatory inputs from both nerves. This input pattern was frequently seen in VS neurons. Approximately half of the convergent VS neurons descended to the spinal cord through the lateral vestibulospinal tract. The remaining half and all the convergent VOS neurons descended to the spinal cord through the medial vestibulospinal tract. Most of the convergent neurons were located in the lateral nucleus or descending nucleus.

Convergence of posterior semicircular canal and saccular inputs in single vestibular nuclei neurons in cats.

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