Abstract:. Gain changes in the vestibuloocular reflex (VOR) during visual-vestibular mismatch stimulation serve as a model system for motor learning. The cerebellar flocculus and its target neurons in the brain stem (FTN) are candidates for the storage of these novel VOR gains. We have recently studied the changes in vertical flocculus Purkinje cells after chronic VOR motor learning. Recently we recorded Y neurons (a vertical type of FTNs) after chronic VOR motor learning and compared these records with vertical floccu… Show more
“…Thus at least a subset of type B cells that have been described in in vitro studies are likely to correspond to the floccular target/eye-head neurons that have previously been described in in vivo studies and have been shown to play an important role in visually induced VOR motor learning in primates (Blazquez et al 2006;Lisberger et al 1994). In alert guinea pig, most VN neurons exhibit a decrease in their head sensitivity in an acute VOR training paradigm (Serafin et al 1999), and recent studies further suggest longer-term changes at the same level underlie memory consolidation required for chronic changes in VOR gain (Kassardjian et al 2005;Shutoh et al 2006).…”
Section: Bridging the Gap: The Relation Between In Vivo And In Vitro mentioning
As a result of the availability of genetic mutant strains and development of noninvasive eye movements recording techniques, the mouse stands as a very interesting model for bridging the gap among behavioral responses, neuronal response dynamics studied in vivo, and cellular mechanisms investigated in vitro. Here we characterized the responses of individual neurons in the mouse vestibular nuclei during vestibular (horizontal whole body rotations) and full field visual stimulation. The majority of neurons (∼2/3) were sensitive to vestibular stimulation but not to eye movements. During the vestibular-ocular reflex (VOR), these neurons discharged in a manner comparable to the “vestibular only” (VO) neurons that have been previously described in primates. The remaining neurons [eye-movement-sensitive (ES) neurons] encoded both head-velocity and eye-position information during the VOR. When vestibular and visual stimulation were applied so that there was sensory conflict, the behavioral gain of the VOR was reduced. In turn, the modulation of sensitivity of VO neurons remained unaffected, whereas that of ES neurons was reduced. ES neurons were also modulated in response to full field visual stimulation that evoked the optokinetic reflex (OKR). Mouse VO neurons, however, unlike their primate counterpart, were not modulated during OKR. Taken together, our results show that the integration of visual and vestibular information in the mouse vestibular nucleus is limited to a subpopulation of neurons which likely supports gaze stabilization for both VOR and OKR.
“…Thus at least a subset of type B cells that have been described in in vitro studies are likely to correspond to the floccular target/eye-head neurons that have previously been described in in vivo studies and have been shown to play an important role in visually induced VOR motor learning in primates (Blazquez et al 2006;Lisberger et al 1994). In alert guinea pig, most VN neurons exhibit a decrease in their head sensitivity in an acute VOR training paradigm (Serafin et al 1999), and recent studies further suggest longer-term changes at the same level underlie memory consolidation required for chronic changes in VOR gain (Kassardjian et al 2005;Shutoh et al 2006).…”
Section: Bridging the Gap: The Relation Between In Vivo And In Vitro mentioning
As a result of the availability of genetic mutant strains and development of noninvasive eye movements recording techniques, the mouse stands as a very interesting model for bridging the gap among behavioral responses, neuronal response dynamics studied in vivo, and cellular mechanisms investigated in vitro. Here we characterized the responses of individual neurons in the mouse vestibular nuclei during vestibular (horizontal whole body rotations) and full field visual stimulation. The majority of neurons (∼2/3) were sensitive to vestibular stimulation but not to eye movements. During the vestibular-ocular reflex (VOR), these neurons discharged in a manner comparable to the “vestibular only” (VO) neurons that have been previously described in primates. The remaining neurons [eye-movement-sensitive (ES) neurons] encoded both head-velocity and eye-position information during the VOR. When vestibular and visual stimulation were applied so that there was sensory conflict, the behavioral gain of the VOR was reduced. In turn, the modulation of sensitivity of VO neurons remained unaffected, whereas that of ES neurons was reduced. ES neurons were also modulated in response to full field visual stimulation that evoked the optokinetic reflex (OKR). Mouse VO neurons, however, unlike their primate counterpart, were not modulated during OKR. Taken together, our results show that the integration of visual and vestibular information in the mouse vestibular nucleus is limited to a subpopulation of neurons which likely supports gaze stabilization for both VOR and OKR.
“…target neurons (FTN) receive input from both the flocculus and primary vestibular afferents and are thought to be important for VOR adaptation (Lisberger 1994;Blazquez et al 2006). The modulation changes of FTN neurons have corresponded with changes in the VOR gain induced by training with visual-vestibular conflict (Lisberger et al 1994a, b;Galiana and Green 1998), so they seem to be a likely main contributor to unilateral VOR adaptation.…”
A recent study showed that the angular vestibuloocular reflex (VOR) can be better adaptively increased using an incremental retinal image velocity error signal compared with a conventional constant large velocity-gain demand (×2). This finding has important implications for vestibular rehabilitation that seeks to improve the VOR response after injury. However, a large portion of vestibular patients have unilateral vestibular hypofunction, and training that raises their VOR response during rotations to both the ipsilesional and contralesional side is not usually ideal. We sought to determine if the vestibular response to one side could selectively be increased without affecting the contralateral response. We tested nine subjects with normal vestibular function. Using the scleral search coil and head impulse techniques, we measured the active and passive VOR gain (eye velocity / head velocity) before and after unilateral incremental VOR adaptation training, consisting of self-generated (active) head impulses, which lasted ∼15 min. The head impulses consisted of rapid, horizontal head rotations with peak-amplitude 15 o , peak-velocity 150 o /s and peak-acceleration 3,000 o /s 2 . The VOR gain towards the adapting side increased after training from 0.92±0.18 to 1.11±0.22 (+22.7± 20.2 %) during active head impulses and from 0.91± 0.15 to 1.01±0.17 (+11.3±7.5 %) during passive head impulses. During active impulses, the VOR gain towards the non-adapting side also increased by ∼8 %, though this increase was ∼70 % less than to the adapting side. A similar increase did not occur during passive impulses. This study shows that unilateral vestibular adaptation is possible in humans with a normal VOR; unilateral incremental VOR adaptation may have a role in vestibular rehabilitation. The increase in passive VOR gain after active head impulse adaptation suggests that the training effect is robust.
“…Prior studies have provided a plausible and testable hypothesis for the neural circuits that mediate the VOR and the sites of cellular changes that cause learning in the VOR (Blazquez et al 2006;Lisberger 1994). The basic circuit is illustrated in Fig.…”
Ramachandran R, Lisberger SG. Neural substrate of modified and unmodified pathways for learning in monkey vestibuloocular reflex. J Neurophysiol 100: 1868 -1878, 2008. First published July 30, 2008 doi:10.1152/jn.90498.2008. To understand how the brain learns, we need to identify the full neural circuit for a behavior; characterize how neural responses in the circuit change during behavioral learning; and understand the nature, location, and control of the cellular changes that are responsible for learning. This goal seems attainable for the vestibuloocular reflex (VOR), where the neural circuit basis for learning is already partially understood. The current hypothesis for VOR learning postulates cellular changes in the cerebellar cortex and the vestibular nucleus. It suggests that the brain stem contains two parallel pathways that have been modeled on the basis of extensive biological data as unmodified and modified VOR pathways with frequency-dependent internal gains and different time delays. We now show a correspondence between the responses of different groups of neurons in the vestibular nucleus and the signals emanating from the two pathways in the model. Floccular target neurons (FTNs) and position-vestibular-pause neurons (PVPs) were identified by their discharge during eye movements and by the presence or absence of inhibition by floccular stimulation. FTNs had response gains and phases that coincided with predictions for pathways that are modified in association with learning, whereas PVPs had responses in agreement with predictions for the unmodified pathways. The quantitative agreement of prior model predictions and new data supports the identity of FTNs and PVPs as brain stem interneurons in the modified and unmodified VOR pathways. Other aspects of the data make predictions about how vestibular inputs are transformed as they pass through the two pathways.
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