Interception of vertically approaching objects: temporal recruitment of the internal model of gravity and contribution of optical information.

Introduction: Recent views posit that precise control of the interceptive timing can be achieved by combining on-line processing of visual information with predictions based on prior experience. Indeed, for interception of free-falling objects under gravity's effects, experimental evidence shows that time-to-contact predictions can be derived from an internal gravity representation in the vestibular cortex. However, whether the internal gravity model is fully engaged at the target motion outset or reinforced by visual motion processing at later stages of motion is not yet clear. Moreover, there is no conclusive evidence about the relative contribution of internalized gravity and optical information in determining the time-to-contact estimates. Methods: We sought to gain insight on this issue by asking 32 participants to intercept free falling objects approaching directly from above in virtual reality. Object motion had durations comprised between 800 and 1100 ms and it could be either congruent with gravity (1 g accelerated motion) or not (constant velocity or -1 g decelerated motion). We analyzed accuracy and precision of the interceptive responses, and fitted them to Bayesian regression models, which included predictors related to the recruitment of a priori gravity information at different times during the target motion, as well as based on available optical information. Results: Consistent with the use of internalized gravity information, interception accuracy and precision were significantly higher with 1 g motion. Moreover, Bayesian regression indicated that interceptive responses were predicted very closely by assuming engagement of the gravity prior 450 ms after the motion onset, and that adding a predictor related to on-line processing of optical information improved only slightly the model predictive power. Discussion: Thus, engagement of a priori gravity information depended critically on the processing of the first 450 ms of visual motion information, exerting a predominant influence on the interceptive timing, compared to continuously available optical information. Finally, these results may support a parallel processing scheme for the control of interceptive timing.

Rhythmic tapping to a moving beat motion kinematics overrules natural gravity.

Beat induction is the cognitive ability that allows humans to listen to a regular pulse in music and move in synchrony with it. Although auditory rhythmic cues induce more consistent synchronization than flashing visual metronomes, this auditory-visual asymmetry can be canceled by visual moving stimuli. Here, we investigated whether the naturalness of visual motion or its kinematics could provide a synchronization advantage over flashing metronomes. Subjects were asked to tap in sync with visual metronomes defined by vertically accelerating/decelerating motion, either congruent or not with natural gravity; horizontally accelerating/decelerating motion; or flashing stimuli. We found that motion kinematics was the predominant factor determining rhythm synchronization, as accelerating moving metronomes in any cardinal direction produced more precise and predictive tapping than decelerating or flashing conditions. Our results support the notion that accelerating visual metronomes convey a strong sense of beat, as seen in the cueing movements of an orchestra director.

A case report of agoraphobia following right parietal lobe surgery: changes in functional and structural connectivities of the multimodal vestibular network.

Agoraphobia is a visuo-vestibular-spatial disorder that may involve dysfunction of the vestibular network, which includes the insular and limbic cortex. We sought to study the neural correlates of this disorder in an individual who developed agoraphobia after surgical removal of a high-grade glioma located in the right parietal lobe, by assessing pre- and post-surgery connectivities in the vestibular network. The patient underwent surgical resection of the glioma located within the right supramarginal gyrus. The resection interested also portions of the superior and inferior parietal lobe. Structural and functional connectivities were assessed through magnetic resonance imaging before and 5 and 7 months after surgery. Connectivity analyses focused on a network comprising 142 spherical regions of interest (4 mm radius) associated with the vestibular cortex: 77 in the left and 65 in the right hemisphere (excluding lesioned regions). Tractography for diffusion-weighted structural data and correlation between time series for functional resting-state data were calculated for each pair of regions in order to build weighted connectivity matrices. Graph theory was applied to assess post-surgery changes in network measures, such as strength, clustering coefficient, and local efficiency. Structural connectomes after surgery showed a decrease of strength in the preserved ventral portion of the supramarginal gyrus (PFcm) and in a high order visual motion area in the right middle temporal gyrus (37dl), and decrease of the clustering coefficient and of the local efficiency in several areas of the limbic, insular cortex, parietal and frontal cortex, indicating general disconnection of the vestibular network. Functional connectivity analysis showed both a decrease in connectivity metrics, mainly in high-order visual areas and in the parietal cortex, and an increase in connectivity metrics, mainly in the precuneus, parietal and frontal opercula, limbic, and insular cortex. This post-surgery reorganization of the vestibular network is compatible with altered processing of visuo-vestibular-spatial information, yielding agoraphobia symptoms. Specifically, post-surgical functional increases of clustering coefficient and local efficiency in the anterior insula and in the cingulate cortex might indicate a more predominant role of these areas within the vestibular network, which could be predictive of the fear and avoiding behavior characterizing agoraphobia.

Watching the Effects of Gravity. Vestibular Cortex and the Neural Representation of "Visual" Gravity.

Gravity is a physical constraint all terrestrial species have adapted to through evolution. Indeed, gravity effects are taken into account in many forms of interaction with the environment, from the seemingly simple task of maintaining balance to the complex motor skills performed by athletes and dancers. Graviceptors, primarily located in the vestibular otolith organs, feed the Central Nervous System with information related to the gravity acceleration vector. This information is integrated with signals from semicircular canals, vision, and proprioception in an ensemble of interconnected brain areas, including the vestibular nuclei, cerebellum, thalamus, insula, retroinsula, parietal operculum, and temporo-parietal junction, in the so-called vestibular network. Classical views consider this stage of multisensory integration as instrumental to sort out conflicting and/or ambiguous information from the incoming sensory signals. However, there is compelling evidence that it also contributes to an internal representation of gravity effects based on prior experience with the environment. This a priori knowledge could be engaged by various types of information, including sensory signals like the visual ones, which lack a direct correspondence with physical gravity. Indeed, the retinal accelerations elicited by gravitational motion in a visual scene are not invariant, but scale with viewing distance. Moreover, the "visual" gravity vector may not be aligned with physical gravity, as when we watch a scene on a tilted monitor or in weightlessness. This review will discuss experimental evidence from behavioral, neuroimaging (connectomics, fMRI, TMS), and patients' studies, supporting the idea that the internal model estimating the effects of gravity on visual objects is constructed by transforming the vestibular estimates of physical gravity, which are computed in the brainstem and cerebellum, into internalized estimates of virtual gravity, stored in the vestibular cortex. The integration of the internal model of gravity with visual and non-visual signals would take place at multiple levels in the cortex and might involve recurrent connections between early visual areas engaged in the analysis of spatio-temporal features of the visual stimuli and higher visual areas in temporo-parietal-insular regions.

Structural connectome and connectivity lateralization of the multimodal vestibular cortical network.

Unlike other sensory systems, the structural connectivity patterns of the human vestibular cortex remain a matter of debate. Based on their functional properties and hypothesized centrality within the vestibular network, the 'core' cortical regions of this network are thought to be areas in the posterior peri-sylvian cortex, in particular the retro-insula (previously named the posterior insular cortex-PIC), and the subregion OP2 of the parietal operculum. To study the vestibular network, structural connectivity matrices from n=974 healthy individuals drawn from the public Human Connectome Project (HCP) repository were estimated using multi-shell diffusion-weighted data followed by probabilistic tractography and spherical-deconvolution informed filtering of tractograms in combination with subject-specific grey-matter parcellations. Weighted graph-theoretical measures, modularity, and 'hubness' of the multimodal vestibular network were then estimated, and a structural lateralization index was defined in order to assess the difference in fiber density of homonym regions in the right and left hemisphere. Differences in connectivity patterns between OP2 and PIC were also estimated. We found that the bilateral intraparietal sulcus, PIC, and to a lesser degree OP2, are key 'hub' regions within the multimodal vestibular network. PIC and OP2 structural connectivity patterns were lateralized to the left hemisphere, while structural connectivity patterns of the posterior peri-sylvian supramarginal and superior temporal gyri were lateralized to the right hemisphere. These lateralization patterns were independent of handedness. We also found that the structural connectivity pattern of PIC is consistent with a key role of PIC in visuo-vestibular processing and that the structural connectivity pattern of OP2 is consistent with integration of mainly vestibular somato-sensory and motor information. These results suggest an analogy between PIC and the simian visual posterior sylvian (VPS) area and OP2 and the simian parieto-insular vestibular cortex (PIVC). Overall, these findings may provide novel insights to the current models of vestibular function, as well as to the understanding of the complexity and lateralized signs of vestibular syndromes.

The role of cortical areas hMT/V5+ and TPJ on the magnitude of representational momentum and representational gravity: a transcranial magnetic stimulation study.

The perceived vanishing location of a moving target is systematically displaced forward, in the direction of motion-representational momentum-, and downward, in the direction of gravity-representational gravity. Despite a wealth of research on the factors that modulate these phenomena, little is known regarding their neurophysiological substrates. The present experiment aims to explore which role is played by cortical areas hMT/V5+, linked to the processing of visual motion, and TPJ, thought to support the functioning of an internal model of gravity, in modulating both effects. Participants were required to perform a standard spatial localization task while the activity of the right hMT/V5+ or TPJ sites was selectively disrupted with an offline continuous theta-burst stimulation (cTBS) protocol, interspersed with control blocks with no stimulation. Eye movements were recorded during all spatial localizations. Results revealed an increase in representational gravity contingent on the disruption of the activity of hMT/V5+ and, conversely, some evidence suggested a bigger representational momentum when TPJ was stimulated. Furthermore, stimulation of hMT/V5+ led to a decreased ocular overshoot and to a time-dependent downward drift of gaze location. These outcomes suggest that a reciprocal balance between perceived kinematics and anticipated dynamics might modulate these spatial localization responses, compatible with a push-pull mechanism.

Ocular tracking of occluded ballistic trajectories: Effects of visual context and of target law of motion.

In tracking a moving target, the visual context may provide cues for an observer to interpret the causal nature of the target motion and extract features to which the visual system is weakly sensitive, such as target acceleration. This information could be critical when vision of the target is temporarily impeded, requiring visual motion extrapolation processes. Here we investigated how visual context influences ocular tracking of motion either congruent or not with natural gravity. To this end, 28 subjects tracked computer-simulated ballistic trajectories either perturbed in the descending segment with altered gravity effects (0g/2g) or retaining natural-like motion (1g). Shortly after the perturbation (550 ms), targets disappeared for either 450 or 650 ms and became visible again until landing. Target motion occurred with either quasi-realistic pictorial cues or a uniform background, presented in counterbalanced order. We analyzed saccadic and pursuit movements after 0g and 2g target-motion perturbations and for corresponding intervals of unperturbed 1g trajectories, as well as after corresponding occlusions. Moreover, we considered the eye-to-target distance at target reappearance. Tracking parameters differed significantly between scenarios: With a neutral background, eye movements did not depend consistently on target motion, whereas with pictorial background they showed significant dependence, denoting better tracking of accelerated targets. These results suggest that oculomotor control is tuned to realistic properties of the visual scene.

Temporomandibular joint reconstruction with stock and custom-made devices: Indications and results of a 14-year experience.

PURPOSE: Temporomandibular joint (TMJ) reconstruction with alloplastic implants is a reliable, safe and effective treatment option for selected debilitating pathologies of the TMJ. This study retrospectively analyzes our 14-year experience in total alloplastic reconstruction of the TMJ using stock and custom Biomet prostheses. Indications, results and rationale for the choice of stock and custom-made devices are discussed. MATERIAL AND METHODS: Patients were enrolled in the study who underwent single-stage alloplastic total joint replacement from January 2000 to October 2014. The subjective and objective variables were as follows: TMJ pain, diet, jaw function, maximum interincisal opening (MIO), quality of life and occlusion. The minimum follow-up was 12 months. RESULTS: A total of 38 patients (55 joints) met the inclusion criteria and were enrolled in the study. Of the patients, 25 underwent Biomet total joint reconstruction system with stock prosthesis, 12 patients underwent total joint reconstruction system with custom made (patient matched) prosthesis, and 1 patient underwent bilateral total joint reconstruction using stock system on one side and custom system on the other side. The following adverse events and complications were recorded: bleeding, 2 cases; malocclusion, 1 case; postoperative infection with prosthesis removal, 1 case; heterotopic bone formation, 1 case; and contralateral TMJ overload in unilateral cases, 1 case. The occlusion was habitual unchanged in 29 of 38 cases. In 1 patient occlusion worsened with less stable functional contact. The patient refused postoperative orthodontic treatment. In 8 patients, a concomitant orthognatic procedure was planned in order to improve the occlusion. In all these patients, the occlusion improved. Quality of life and MIO relevantly improved in all cases. CONCLUSIONS: This study supports the use of total joint reconstruction for end-stage TMJ disease. Both stock and custom implants allow consistent results, but there are precise indications for the use of custom implants.

Differential contributions to the interception of occluded ballistic trajectories by the temporoparietal junction, area hMT/V5+, and the intraparietal cortex.

The ability to catch objects when transiently occluded from view suggests their motion can be extrapolated. Intraparietal cortex (IPS) plays a major role in this process along with other brain structures, depending on the task. For example, interception of objects under Earth's gravity effects may depend on time-to-contact predictions derived from integration of visual signals processed by hMT/V5+ with a priori knowledge of gravity residing in the temporoparietal junction (TPJ). To investigate this issue further, we disrupted TPJ, hMT/V5+, and IPS activities with transcranial magnetic stimulation (TMS) while subjects intercepted computer-simulated projectile trajectories perturbed randomly with either hypo- or hypergravity effects. In experiment 1, trajectories were occluded either 750 or 1,250 ms before landing. Three subject groups underwent triple-pulse TMS (tpTMS, 3 pulses at 10 Hz) on one target area (TPJ | hMT/V5+ | IPS) and on the vertex (control site), timed at either trajectory perturbation or occlusion. In experiment 2, trajectories were entirely visible and participants received tpTMS on TPJ and hMT/V5+ with same timing as experiment 1 tpTMS of TPJ, hMT/V5+, and IPS affected differently the interceptive timing. TPJ stimulation affected preferentially responses to 1-g motion, hMT/V5+ all response types, and IPS stimulation induced opposite effects on 0-g and 2-g responses, being ineffective on 1-g responses. Only IPS stimulation was effective when applied after target disappearance, implying this area might elaborate memory representations of occluded target motion. Results are compatible with the idea that IPS, TPJ, and hMT/V5+ contribute to distinct aspects of visual motion extrapolation, perhaps through parallel processing.NEW & NOTEWORTHY Visual extrapolation represents a potential neural solution to afford motor interactions with the environment in the face of missing information. We investigated relative contributions by temporoparietal junction (TPJ), hMT/V5+, and intraparietal cortex (IPS), cortical areas potentially involved in these processes. Parallel organization of visual extrapolation processes emerged with respect to the target's motion causal nature: TPJ was primarily involved for visual motion congruent with gravity effects, IPS for arbitrary visual motion, whereas hMT/V5+ contributed at earlier processing stages.

Gravity in the Brain as a Reference for Space and Time Perception.

Moving and interacting with the environment require a reference for orientation and a scale for calibration in space and time. There is a wide variety of environmental clues and calibrated frames at different locales, but the reference of gravity is ubiquitous on Earth. The pull of gravity on static objects provides a plummet which, together with the horizontal plane, defines a three-dimensional Cartesian frame for visual images. On the other hand, the gravitational acceleration of falling objects can provide a time-stamp on events, because the motion duration of an object accelerated by gravity over a given path is fixed. Indeed, since ancient times, man has been using plumb bobs for spatial surveying, and water clocks or pendulum clocks for time keeping. Here we review behavioral evidence in favor of the hypothesis that the brain is endowed with mechanisms that exploit the presence of gravity to estimate the spatial orientation and the passage of time. Several visual and non-visual (vestibular, haptic, visceral) cues are merged to estimate the orientation of the visual vertical. However, the relative weight of each cue is not fixed, but depends on the specific task. Next, we show that an internal model of the effects of gravity is combined with multisensory signals to time the interception of falling objects, to time the passage through spatial landmarks during virtual navigation, to assess the duration of a gravitational motion, and to judge the naturalness of periodic motion under gravity.

Filling gaps in visual motion for target capture.

A remarkable challenge our brain must face constantly when interacting with the environment is represented by ambiguous and, at times, even missing sensory information. This is particularly compelling for visual information, being the main sensory system we rely upon to gather cues about the external world. It is not uncommon, for example, that objects catching our attention may disappear temporarily from view, occluded by visual obstacles in the foreground. Nevertheless, we are often able to keep our gaze on them throughout the occlusion or even catch them on the fly in the face of the transient lack of visual motion information. This implies that the brain can fill the gaps of missing sensory information by extrapolating the object motion through the occlusion. In recent years, much experimental evidence has been accumulated that both perceptual and motor processes exploit visual motion extrapolation mechanisms. Moreover, neurophysiological and neuroimaging studies have identified brain regions potentially involved in the predictive representation of the occluded target motion. Within this framework, ocular pursuit and manual interceptive behavior have proven to be useful experimental models for investigating visual extrapolation mechanisms. Studies in these fields have pointed out that visual motion extrapolation processes depend on manifold information related to short-term memory representations of the target motion before the occlusion, as well as to longer term representations derived from previous experience with the environment. We will review recent oculomotor and manual interception literature to provide up-to-date views on the neurophysiological underpinnings of visual motion extrapolation.

Eye movements and manual interception of ballistic trajectories: effects of law of motion perturbations and occlusions.

Manual interceptions are known to depend critically on integration of visual feedback information and experience-based predictions of the interceptive event. Within this framework, coupling between gaze and limb movements might also contribute to the interceptive outcome, since eye movements afford acquisition of high-resolution visual information. We investigated this issue by analyzing subjects' head-fixed oculomotor behavior during manual interceptions. Subjects moved a mouse cursor to intercept computer-generated ballistic trajectories either congruent with Earth's gravity or perturbed with weightlessness (0 g) or hypergravity (2 g) effects. In separate sessions, trajectories were either fully visible or occluded before interception to enforce visual prediction. Subjects' oculomotor behavior was classified in terms of amounts of time they gazed at different visual targets and of overall number of saccades. Then, by way of multivariate analyses, we assessed the following: (1) whether eye movement patterns depended on targets' laws of motion and occlusions; and (2) whether interceptive performance was related to the oculomotor behavior. First, we found that eye movement patterns depended significantly on targets' laws of motion and occlusion, suggesting predictive mechanisms. Second, subjects coupled differently oculomotor and interceptive behavior depending on whether targets were visible or occluded. With visible targets, subjects made smaller interceptive errors if they gazed longer at the mouse cursor. Instead, with occluded targets, they achieved better performance by increasing the target's tracking accuracy and by avoiding gaze shifts near interception, suggesting that precise ocular tracking provided better trajectory predictions for the interceptive response.

Multisensory integration and internal models for sensing gravity effects in primates.

Gravity is crucial for spatial perception, postural equilibrium, and movement generation. The vestibular apparatus is the main sensory system involved in monitoring gravity. Hair cells in the vestibular maculae respond to gravitoinertial forces, but they cannot distinguish between linear accelerations and changes of head orientation relative to gravity. The brain deals with this sensory ambiguity (which can cause some lethal airplane accidents) by combining several cues with the otolith signals: angular velocity signals provided by the semicircular canals, proprioceptive signals from muscles and tendons, visceral signals related to gravity, and visual signals. In particular, vision provides both static and dynamic signals about body orientation relative to the vertical, but it poorly discriminates arbitrary accelerations of moving objects. However, we are able to visually detect the specific acceleration of gravity since early infancy. This ability depends on the fact that gravity effects are stored in brain regions which integrate visual, vestibular, and neck proprioceptive signals and combine this information with an internal model of gravity effects.

Visual gravitational motion and the vestibular system in humans.

The visual system is poorly sensitive to arbitrary accelerations, but accurately detects the effects of gravity on a target motion. Here we review behavioral and neuroimaging data about the neural mechanisms for dealing with object motion and egomotion under gravity. The results from several experiments show that the visual estimates of a target motion under gravity depend on the combination of a prior of gravity effects with on-line visual signals on target position and velocity. These estimates are affected by vestibular inputs, and are encoded in a visual-vestibular network whose core regions lie within or around the Sylvian fissure, and are represented by the posterior insula/retroinsula/temporo-parietal junction. This network responds both to target motions coherent with gravity and to vestibular caloric stimulation in human fMRI studies. Transient inactivation of the temporo-parietal junction selectively disrupts the interception of targets accelerated by gravity.

Catching what we can't see: manual interception of occluded fly-ball trajectories.

Control of interceptive actions may involve fine interplay between feedback-based and predictive mechanisms. These processes rely heavily on target motion information available when the target is visible. However, short-term visual memory signals as well as implicit knowledge about the environment may also contribute to elaborate a predictive representation of the target trajectory, especially when visual feedback is partially unavailable because other objects occlude the visual target. To determine how different processes and information sources are integrated in the control of the interceptive action, we manipulated a computer-generated visual environment representing a baseball game. Twenty-four subjects intercepted fly-ball trajectories by moving a mouse cursor and by indicating the interception with a button press. In two separate sessions, fly-ball trajectories were either fully visible or occluded for 750, 1000 or 1250 ms before ball landing. Natural ball motion was perturbed during the descending trajectory with effects of either weightlessness (0 g) or increased gravity (2 g) at times such that, for occluded trajectories, 500 ms of perturbed motion were visible before ball disappearance. To examine the contribution of previous visual experience with the perturbed trajectories to the interception of invisible targets, the order of visible and occluded sessions was permuted among subjects. Under these experimental conditions, we showed that, with fully visible targets, subjects combined servo-control and predictive strategies. Instead, when intercepting occluded targets, subjects relied mostly on predictive mechanisms based, however, on different type of information depending on previous visual experience. In fact, subjects without prior experience of the perturbed trajectories showed interceptive errors consistent with predictive estimates of the ball trajectory based on a-priori knowledge of gravity. Conversely, the interceptive responses of subjects previously exposed to fully visible trajectories were compatible with the fact that implicit knowledge of the perturbed motion was also taken into account for the extrapolation of occluded trajectories.

Representation of movement velocity in the rat's interpositus nucleus during passive forelimb movements.

The interpositus nucleus (IN) receives a large amount of sensory information from the limbs and, in turn, elaborates signals for movement control. In this paper, we tried to gather evidence on the possibility that neurons in the IN may elaborate sensory representations of the forelimb kinematics and, particularly, of the movement velocity vector. For this purpose, the forepaw of anesthetized rats was attached to a computer-controlled robot arm displaced passively along two types of trajectories (circular and figure eight), with the limb joints unconstrained. The firing activity of single cells was recorded and related to limb position and the two components of the movement velocity vector, namely, movement speed and direction. By using multiple regression analysis, we found that 12 out of 85 (14%) neurons were modulated by position, 18 out of 85 (21%) neurons were modulated by direction, 24 out of 85 (28%) neurons were modulated by movement speed, and 31 out of 85 (37%) neurons were sensitive to the full movement velocity vector. Most of the neurons modulated only by the speed component of the velocity vector (19 out of 24) were located in the posterior portion of the IN, whereas neurons in the anterior portion were mostly related to both components of the velocity vector. These results suggest that sensory information related to whole-limb movement velocity may be encoded by the IN, indicating also that the posterior interpositus may preferentially represent movement speed.

Processing of limb kinematics in the interpositus nucleus.

Neural representations of limb movement kinematic parameters are common among central nervous system structures involved in motor control, such as the interpositus nucleus of the cerebellum. Much experimental evidence indicates that neurons in the interpositus may encode limb kinematic parameters both during active, voluntary actions and during limb motion imposed passively, which entrains only sensory afferents. With respect to the sensory processing of information related to movement kinematics, we show that interpositus neuronal activity can parse out the directional from the scalar component (i.e., the movement speed) of the velocity vector. Moreover, a differential role for the anterior and posterior portion of interpositus in encoding these parameters emerged from these data, since the activity of the posterior interpositus was specifically associated to changes of movement speed. Limb movement representations in the interpositus nucleus may be instrumental for the control of goal-directed movements such as shaping hand during grasping or precise foot placement during gait. Finally, we discuss the idea that sensory information about the movement kinematics contribute to both feedback and anticipatory processes for limb movement control.

Contributions of the human temporoparietal junction and MT/V5+ to the timing of interception revealed by transcranial magnetic stimulation.

To intercept a fast target at destination, hand movements must be centrally triggered ahead of target arrival to compensate for neuromechanical delays. The role of visual-motion cortical areas is unclear. They likely feed downstream parietofrontal networks with signals reflecting target motion, but do they also contribute internal timing signals to trigger the motor response? We disrupted the activity of human temporoparietal junction (TPJ) and middle temporal area (hMT/V5+) by means of transcranial magnetic stimulation (TMS) while subjects pressed a button to intercept targets accelerated or decelerated in the vertical or horizontal direction. Target speed was randomized, making arrival time unpredictable across trials. We used either repetitive TMS (rTMS) before task execution or double-pulse TMS (dpTMS) during target motion. We found that after rTMS and dpTMS at 100-200 ms from motion onset, but not after dpTMS at 300-400 ms, the button-press responses occurred earlier than in the control, with time shifts independent of target speed. This suggests that activity in TPJ and hMT/V5+ can feed downstream regions not only with visual-motion information, but also with internal timing signals used for interception at destination. Moreover, we found that TMS of hMT/V5+ affected interception of all tested motion types, whereas TMS of TPJ significantly affected only interception of motion coherent with natural gravity. TPJ might specifically gate visual-motion information according to an internal model of the effects of gravity.

Vestibular nuclei and cerebellum put visual gravitational motion in context.

Animal survival in the forest, and human success on the sports field, often depend on the ability to seize a target on the fly. All bodies fall at the same rate in the gravitational field, but the corresponding retinal motion varies with apparent viewing distance. How then does the brain predict time-to-collision under gravity? A perspective context from natural or pictorial settings might afford accurate predictions of gravity's effects via the recovery of an environmental reference from the scene structure. We report that embedding motion in a pictorial scene facilitates interception of gravitational acceleration over unnatural acceleration, whereas a blank scene eliminates such bias. Functional magnetic resonance imaging (fMRI) revealed blood-oxygen-level-dependent correlates of these visual context effects on gravitational motion processing in the vestibular nuclei and posterior cerebellar vermis. Our results suggest an early stage of integration of high-level visual analysis with gravity-related motion information, which may represent the substrate for perceptual constancy of ubiquitous gravitational motion.

Representation of visual gravitational motion in the human vestibular cortex.

How do we perceive the visual motion of objects that are accelerated by gravity? We propose that, because vision is poorly sensitive to accelerations, an internal model that calculates the effects of gravity is derived from graviceptive information, is stored in the vestibular cortex, and is activated by visual motion that appears to be coherent with natural gravity. The acceleration of visual targets was manipulated while brain activity was measured using functional magnetic resonance imaging. In agreement with the internal model hypothesis, we found that the vestibular network was selectively engaged when acceleration was consistent with natural gravity. These findings demonstrate that predictive mechanisms of physical laws of motion are represented in the human brain.