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We try to deploy the retinal fovea to optimally scrutinize an object of interest by directing our eyes to it. The horizontal and vertical components of eye positions acquired by goal-directed saccades are determined by the object's location. However, the eccentric eye positions also involve a torsional component, which according to Donder's law is fully determined by the two-dimensional (2D) eye position acquired. According to von Helmholtz, knowledge of the amount of torsion provided by Listing's law, an extension of Donder's law, alleviates the perceptual interpretation of the image tilt that changes with 2D eye position, a view supported by psychophysical experiments he pioneered. We address the question of where and how Listing's law is implemented in the visual system and we show that neurons in monkey area V1 use knowledge of eye torsion to compensate the image tilt associated with specific eye positions as set by Listing's law., Competing Interests: The authors declare no competing interest.

Recent studies have suggested that microsaccades, the small amplitude saccades made during fixation, are precisely controlled. Two lines of evidence suggest that the cerebellum plays a key role not only in improving the accuracy of macrosaccades but also of microsaccades. First, lesions of the fastigial oculomotor regions (FOR) cause horizontal dysmetria of both micro- and macrosaccades. Secondly, our previous work on Purkinje cell simple spikes in the oculomotor vermis (OV) has established qualitatively similar response preferences for these two groups of saccades. In this work, we investigated the control signals for micro- and macrosaccades in the FOR, the target of OV Purkinje cell axons. We found that the same FOR neurons discharged for micro- and macrosaccades. For both groups of saccades, FOR neurons exhibited very similar dependencies of their discharge strength on direction and amplitude and very similar burst onset time differences for ipsi- and contraversive saccades and, in both, response duration reflected saccade duration, at least at the population level. An intriguing characteristic of microsaccade-related responses is that immediate pre-saccadic firing rates decreased with distance to the target center, a pattern that strikingly parallels the eye position dependency of both microsaccade metrics and frequency, which may suggest a potential neural mechanism underlying the role of FOR in fixation. Irrespective of this specific consideration, our study supports the view that microsaccades and macrosaccades share the same cerebellar circuitry and, in general, further strengthens the notion of a microsaccade-macrosaccade continuum., (© 2016 Federation of European Neuroscience Societies and John Wiley & Sons Ltd.)

The pontine nuclei (PN) are the major intermediary elements in the corticopontocerebellar pathway. Here we asked if the PN may help to adapt the spatial reference frames used by cerebrocortical neurons involved in the sensory guidance of movement to a format potentially more appropriate for the cerebellum. To this end, we studied movement-related neurons in the dorsal PN (DPN) of monkeys, most probably projecting to the cerebellum, executing fixed vector saccades or, alternatively, fixed vector hand reaches from different starting positions. The 83 task-related neurons considered fired movement-related bursts before saccades (saccade-related) or before hand movements (hand movement-related). About 40% of the SR neurons were "oculocentric," whereas the others were modulated by eye starting position. A third of the HMR neurons encoded hand reaches in hand-centered coordinates, whereas the remainder exhibited different types of dependencies on starting positions, reminiscent in general of cortical responses. All in all, pontine reference frames for the sensory guidance of movement seem to be very similar to those in cortex. Specifically, the frequency of orbital position gain fields of SR neurons is identical in the DPN and in one of their major cortical inputs, lateral intraparietal area (LIP).

How does the nervous system guide the muscular periphery during the acquisition of a new motor skill? This is a fundamental question for researchers trying to understand the neural basis of motor learning. Recent advances in studying a valuable example of short-term motor learning, namely the adaptation of saccadic eye movements, have revealed neuronal processes in the cerebellum that underlie the unfolding of the learned behavior. In this review, we describe the latest findings from electrophysiology studies of saccadic adaptation and how they can generalize to more elaborate examples of cerebellum-dependent adaptation of movements. We focus our discussion on the plastic changes that are observed in the firing properties of Purkinje cells during the acquisition of the wanted motor response and describe how the altered activity of these neurons modifies the dynamics of the cerebellar microcircuitry. We finally demonstrate how such task-related modifications in the cerebellum are appropriate to fine-tune extracerebellar pre-motor structures and induce the learned behavior., (© 2011 The Authors. European Journal of Neuroscience © 2011 Federation of European Neuroscience Societies and Blackwell Publishing Ltd.)

Converging experimental evidence indicates that mirror neurons in the monkey premotor area F5 encode the goals of observed motor acts [1-3]. However, it is unknown whether they also contribute to encoding the perspective from which the motor acts of others are seen. In order to address this issue, we recorded the visual responses of mirror neurons of monkey area F5 by using a novel experimental paradigm based on the presentation of movies showing grasping motor acts from different visual perspectives. We found that the majority of the tested mirror neurons (74%) exhibited view-dependent activity with responses tuned to specific points of view. A minority of the tested mirror neurons (26%) exhibited view-independent responses. We conclude that view-independent mirror neurons encode action goals irrespective of the details of the observed motor acts, whereas the view-dependent ones might either form an intermediate step in the formation of view independence or contribute to a modulation of view-dependent representations in higher-level visual areas, potentially linking the goals of observed motor acts with their pictorial aspects., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Prevailing concepts on the control of goal-directed hand movements (HM) have focused on a network of cortical areas whose early parieto-occipital stages are thought to extract and integrate information on target and hand location, involving subsequent stages in frontal cortex forming and executing movement plans. The substantial experimental results supporting this "cortical network" concept for hand movements notwithstanding, the concept clearly needs refinement to account for the surprisingly mild HM disturbances resulting from disconnecting the parieto-occipital from the frontal stages of the network. Clinical observations have suggested the cerebropontocerebellar projection as an alternative pathway for the sensory guidance of HM. As a first step in assessing its role, we explored the pontine nuclei (PN) of rhesus monkeys, trained to make goal-directed hand and eye movements guided by spatial memory. We were indeed able to delineate a distinct cluster of neurons in the rostrodorsal PN, activated by the preparation and the execution of hand reaches, close to but distinct from the region in which saccade-related neurons may be observed. The movement-related activity of HM-related neurons starts earlier than that of saccade-related neurons and both neuron types are usually effector specific, i.e., they respond only to the movement of the preferred effector. This is also the case when motor synergies involving both effectors are executed. Our findings support the notion of a distinct precerebellar, pontine visuomotor channel for hand reaches that is anatomically and functionally largely separated from the one serving eye movements.

Actions performed by others may have different relevance for the observer, and thus lead to different behavioral responses, depending on the regions of space in which they are executed. We found that in rhesus monkeys, the premotor cortex neurons activated by both the execution and the observation of motor acts (mirror neurons) are differentially modulated by the location in space of the observed motor acts relative to the monkey, with about half of them preferring either the monkey's peripersonal or extrapersonal space. A portion of these spatially selective mirror neurons encode space according to a metric representation, whereas other neurons encode space in operational terms, changing their properties according to the possibility that the monkey will interact with the object. These results suggest that a set of mirror neurons encodes the observed motor acts not only for action understanding, but also to analyze such acts in terms of features that are relevant to generating appropriate behaviors.

We are usually unaware of retinal image motion resulting from our own movement. For instance, during slow-tracking eye movements the world around us remains perceptually stable despite the retinal image slip induced by the eye movement. It is commonly held that this example of perceptual invariance is achieved by subtracting an internal reference signal, reflecting the eye movement, from the retinal motion signal. If the two cancel each other, visual objects, which do not move, will also be perceived as non-moving. If, however, the reference signal is too small or too large, a false eye movement-induced motion of the external world, the Filehne illusion, will be perceived. We have exploited our ability to manipulate the size of the reference signal in an attempt to identify neurons in the visual cortex of monkeys, influenced by the percept of self-induced visual motion or the reference signal rather than the retinal motion signal. We report here that such 'percept-related' neurons can already be found in the primary visual cortex area, although few in numbers. They become more frequent in areas middle temporal and medial superior temporal in the superior temporal sulcus, and comprise almost 50% of all neurons in area visual posterior sylvian (VPS) in the posterior part of the lateral sulcus. In summary, our findings suggest that our ability to perceive a visual world, which is stable despite self-motion, is based on a neuronal network, which culminates in the VPS located in the lateral sulcus below the classical dorsal stream of visual processing.

The motion areas of posterior parietal cortex extract information on visual motion for perception as well as for the guidance of movement. It is usually assumed that neurons in posterior parietal cortex represent visual motion relative to the retina. Current models describing action guided by moving objects work successfully based on this assumption. However, here we show that the pursuit-related responses of a distinct group of neurons in area MST of monkeys are at odds with this view. Rather than signaling object image motion on the retina, they represent object motion in world-centered coordinates. This representation may simplify the coordination of object-directed action and ego motion-invariant visual perception.

The primate dorsolateral pontine nucleus (DLPN) is a key link in a cerebro-cerebellar pathway for smooth pursuit eye movements, a pathway assumed to be anatomically segregated from tegmental circuits subserving saccades. However, the existence of afferents from several cerebrocortical and subcortical centres for saccades suggests that the DLPN and neighbouring parts of the dorsal pontine nuclei (DPN) might contribute to saccades as well. In order to test this hypothesis, we recorded from the DPN of two monkeys trained to perform smooth pursuit eye movements as well as visually and memory-guided saccades. Out of 281 neurons isolated from the DPN, 138 were responsive in oculomotor tasks. Forty-five were exclusively activated in saccade paradigms, 68 exclusively by smooth pursuit and 25 neurons showed responses in both. Pursuit-related responses reflected sensitivity to eye position, velocity or combinations of velocity and position with minor contributions of acceleration in many cases. When tested in the memory-guided saccades paradigm, 65 out of 70 neurons activated in saccade paradigms showed significant saccade-related bursts and 20 significant activity in the memory period. Our finding of saccade-related activity in the DPN in conjunction with the existence of strong anatomical input from saccade-related cerebrocortical areas suggests that the DPN serves as a precerebellar relay for both pursuit and saccade-related information originating from cerebral cortex, in addition to the classical tecto-tegmental circuitry for saccades.

The analysis of a peripheral visual location can be improved in two ways: either by orienting one's gaze (usually by making a foveating saccade) or by 'covertly' shifting one's attention to the peripheral location without making an eye movement. The premotor theory of attention holds that saccades and spatial shifts of attention share a common functional module with a distinct neuronal basis. Using single-unit recording from the brains of trained rhesus monkeys, we investigated whether the superior colliculus, the major subcortical center for the control of saccades, is part of this shared network for attention and saccades. Here we show that a distinct type of neuron in the intermediate layer of the superior colliculus, the visuomotor neuron, which is known to be centrally involved in the preparation of saccades, is also active during covert shifts of attention.

Because smooth-pursuit eye movements (SPEM) can be executed only in the presence of a moving target, it has been difficult to attribute the neuronal activity observed during the execution of these eye movements to either sensory processing or to motor preparation or execution. Previously, we showed that rhesus monkeys can be trained to perform SPEM directed toward an "imaginary" target defined by visual cues confined to the periphery of the visual field. The pursuit of an "imaginary" target provides the opportunity to elicit SPEM without stimulating visual receptive fields confined to the center of the visual field. Here, we report that a subset of neurons [85 "imaginary" visual tracking (iVT)-neurons] in area MST of 3 rhesus monkeys were identically activated during pursuit of a conventional, foveal dot target and the "imaginary" target. Because iVT-neurons did not respond to the presentation of a moving "imaginary" target during fixation of a stationary dot, we are able to exclude that responses to pursuit of the "imaginary" target were artifacts of stimulation of the visual field periphery. Neurons recorded from the representation of the central parts of the visual field in neighboring area MT, usually vigorously discharging during pursuit of foveal targets, in no case responded to pursuit of the "imaginary" target. This dissociation between MT and MST neurons supports the view that pursuit responses of MT neurons are the result of target image motion, whereas those of iVT-neurons in area MST reflect an eye movement-related signal that is nonretinal in origin. iVT-neurons fell into two groups, depending on the properties of the eye movement-related signal. Whereas most of them (71%) encoded eye velocity, a minority showed responses determined by eye position, irrespective of whether eye position was changed by smooth pursuit or by saccades. Only the former group exhibited responses that led the eye movement, which is a prerequisite for a causal role in the generation of SPEM.

The deep cerebellar nuclei (DCN) constitute the major structures by which the cerebellum forwards its output to the rest of the brain. Although the connectivity of the DCN has been well studied, little is known about the interface-the neurons' soma and dendrites-between the DCN's inputs and outputs. We therefore decided to analyze the neurons' somatic and dendritic morphology by applying a multivariate approach (principal component analysis; PCA), in order to define morphological groups possibly related to distinct positions in the nuclear microcircuitry. The PCA was based on intracellularly stained neurons from the rat's lateral DCN and on 19 parameters that described the neurons' morphology. The PCA yielded two principal components that accounted for 46% of the variance. The first component, correlated with soma size, separated the majority of neurons (type I) from a population of small neurons (type II). The second component showed negative correlation with larger cells with more numerous primary dendrites and a more multipolar appearance (type Ia) and positive correlation with smaller neurons with asymmetric dendritic fields and tufted dendrites (type Ib). The preponderance of small somata in our type Ib neurons suggests that these neurons probably correspond to the inferior olive projection neurons. In summary, our results are in agreement with previous classifications, which distinguished projection neurons (type I) from local neurons (type II); furthermore, our results point to a hitherto undescribed dendritic morphological difference in the projection neurons. The latter may be important for understanding the phylogenetic changes seen in the mammalian lateral cerebellar nucleus., (Copyright 2002 Wiley-Liss, Inc.)

Recent studies have shown that neural activity in a well-defined patch in the posterior superior temporal sulcus (the "gaze-following patch," GFP) of the primate brain is strongly modulated when the other's gaze attracts the observer's attention to locations/ objects, the other is looking at. Changes of the mean discharge rate of neurons in the monkey GFP indicate that they are involved in two distinct computations: the allocation of spatial attention guided by the other's gaze vector and the suppression of gaze following if inappropriate in a given situation. Here, we asked if and how the discharge variability of neurons in the GFP is related to the task and if it carries information on behavioral performance. To this end, we calculated the Fano factor as a measure of across-trial discharge variability as a function of time. Our results show that all neurons exhibiting a task-related discharge- rate modulation also exhibit a stimulus onset-dependent drop in the Fano factor. Furthermore, the amplitude of the Fano factor reduction is modulated by task condition and the neuron's selectivity in this regard. We found that these effects are directly related to the monkeys' behavioral performance in that the Fano factor is predictive about upcoming correct or wrong decisions. Our results indicate that neuronal discharge variability as gauged by the Fano factor, hitherto primarily studied in the context of visual perception or motor control, is an informative measure also in studies of the neural underpinnings of complex social behavior. [ABSTRACT FROM AUTHOR]

The cerebellum allows us to rapidly adjust motor behavior to the needs of the situation. It is commonly assumed that cerebellum-based motor learning is guided by the difference between the desired and the actual behavior, i.e., by error information. Not only immediate but also future behavior will benefit from an error because it induces lasting changes of parallel fiber synapses on Purkinje cells (PCs), whose output mediates the behavioral adjustments. Olivary climbing fibers, likewise connecting with PCs, are thought to transport information on instant errors needed for the synaptic modification yet not to contribute to error memory. Here, we report work on monkeys tested in a saccadic learning paradigm that challenges this concept. We demonstrate not only a clear complex spikes (CS) signature of the error at the time of its occurrence but also a reverberation of this signature much later, before a new manifestation of the behavior, suitable to improve it. [ABSTRACT FROM AUTHOR]

Responses of primary visual cortex (V1) neurons to stimuli inside the classic receptive field (CRF) can be modulated by stimuli outside the CRF. We recently reported that responses of most V1 neurons to a line in the CRF center are inhibited by large surround-stimuli and that this modulation is stimulus selective. Here we report that a significant proportion of V1 neurons in alert monkeys respond directly to stimuli outside the CRF with very long latency and much reduced selectivity. When surround stimuli are presented alone, three response patterns can be distinguished in 153 single- or multiunits tested: (1) 31.4% have no significant response; (2) 50.3% show excitatory responses that are significantly higher than spontaneous activity. The average latency of these responses is about 145 ms, 2–3 times longer than center responses; (3) 18.3% show suppressed spontaneous activity after stimulus onset. The direct surround responses are found to be only weakly selective for the orientation of contextual lines, and not selective for other contextual patterns tested. While the outburst of responses to stimuli within the CRF is not affected by reducing stimulus duration from 500 ms to 50 ms, late excitatory surround responses are virtually eliminated. We propose that the late excitatory surround responses to extra-CRF stimulation alone are the reflection of feedback from higher cortical areas and may contribute to reduced contextual inhibition of cells in V1. This could play a role in figure-ground segregation. [ABSTRACT FROM AUTHOR]

Focuses on the pontine nuclei (PN) which are intercalated in the cerebro-cerebellar pathway. How the intrinsic properties of PN neurons permit complex control of signal flow to the cerebellum; Variability of neurons in its dendritic morphology; Cortical signals in the PN neurons.

Abstract. The cortical medial superior temporal area (MST) is essential for the normal execution of smooth pursuit eye movements. Many pursuit-related neurons (visual-tracking neurons = VT neurons) in the lateral part of area MST (MST1) are responsive to retinal image slip (r) as well as to eye (e) and head velocity (h) with similar preferred directions (isodirectionality). We show, by running a connectionist network with VT neuron-like elements, that an assembly of MST1-VT neurons is able to reconstruct target motion in world-centered coordinates (t'). When t' is fed into a subsequent model stage, converting t' into gaze velocity (g') with varying contributions of e and h, the overall model is able to account for many of the salient properties of visually guided pursuit including the consequences of MST1 lesions. However, the analysis of the MST1 network also clearly indicates that isodirectionality is not a prerequisite for its performance. The investigation of a second model suggests that isodirectionality indeed does not result from functional but from developmental constraints. This second model is a connectionist network with hidden units, which similar to MST1-VT neurons receive input from modality specific units encoding retinal slip, eye and head velocity. After training this network to offer t' as output, two subsets of hidden units emerged, one exhibiting isodirectionality, but not the other. Since only isodirectional hidden units contributed to the flow of information, the preponderance of isodirectional MSTI-VT neurons might be the result of developmental pruning, eliminating the second group. [ABSTRACT FROM AUTHOR]

An abstract of the article "Saccade angle modulate correlation between the local field potential and cerebellar Purkinje neuron activity" by Sungho Hong, Mario Negrello, Marc A. Junker, and colleagues is presented.

According to the mirror mechanism the discharge of F5 mirror neurons of a monkey observing another individual performing an action is a motor representation of the observed action that may serve to understand or learn from the action. This hypothesis, if strictly interpreted, requires mirror neurons to exhibit an action tuning that is shared between action observation and execution. Due to insufficient data it remains contentious if this requirement is met. To fill in the gaps, we conducted an experiment in which identical objects had to be manipulated in three different ways in order to serve distinct action goals. Using three methods, including cross-task classification, we found that at most time points F5 mirror neurons did not encode observed actions with the same code underlying action execution. However, in about 20% of neurons there were time periods with a shared code. These time periods formed a distinct cluster and cannot be considered a product of chance. Population classification yielded non-shared coding for observed actions in the whole population, which was at times optimal and consistently better than shared coding in differentially selected subpopulations. These results support the hypothesis of a representation of observed actions based on a strictly defined mirror mechanism only for small subsets of neurons and only under the assumption of time-resolved readout. Considering alternative concepts and recent findings, we propose that during observation mirror neurons represent the process of a goal pursuit from the observer's viewpoint. Whether the observer's goal pursuit, in which the other's action goal becomes the observer's action goal, or the other's goal pursuit is represented remains to be clarified. In any case, it may allow the observer to use expectations associated with a goal pursuit to directly intervene in or learn from another's action. [ABSTRACT FROM AUTHOR]

Purkinje cells (PC), the sole output neurons of the cerebellar cortex, encode sensorimotor information, but how they do it remains a matter of debate. Here we show that PCs use a multiplexed spike code. Synchrony/spike time and firing rate encode different information in behaving monkeys during saccadic eye motion tasks. Using the local field potential (LFP) as a probe of local network activity, we found that infrequent pause spikes, which initiated or terminated intermittent pauses in simple spike trains, provide a temporally reliable signal for eye motion onset, with strong phase-coupling to the β/γ band LFP. Concurrently, regularly firing, non-pause spikes were weakly correlated with the LFP, but were crucial to linear encoding of eye movement kinematics by firing rate. Therefore, PC spike trains can simultaneously convey information necessary to achieve precision in both timing and continuous control of motion. [ABSTRACT FROM AUTHOR]

Abstract: After having been exposed to strong visual motion in one direction, a subsequently presented stationary visual scene seems to move in the opposite direction. This motion aftereffect (MAE) is usually ascribed to short-term functional changes in cortical areas involved in visual motion analysis akin to adaptation. Using magnetoencephalography (MEG), we show increased global field activity due to the MAE which could mostly be explained by a dipole located near the putative location of human area MT+. We further demonstrate that the induced MAE is accompanied by a significant increase in gamma-band activity (GBA) recorded from parietooccipital cortex contralateral to the visual motion stimulus. This gamma oscillation most likely reflects an increase in neuronal response coherence due to decreased inhibition of a group of neurons with similar preferred direction, namely the direction opposite to the adapted one. A second focal GBA response was picked up by the most posterior sensors ipsilateral to the side of the stimulus, reflecting the size of the MAE, whose source could not be reliably located. [Copyright &y& Elsevier]