Neural sensitivity to translational self- and object-motion velocities.

The ability to detect and assess world-relative object-motion is a critical computation performed by the visual system. This computation, however, is greatly complicated by the observer's movements, which generate a global pattern of motion on the observer's retina. How the visual system implements this computation is poorly understood. Since we are potentially able to detect a moving object if its motion differs in velocity (or direction) from the expected optic flow generated by our own motion, here we manipulated the relative motion velocity between the observer and the object within a stationary scene as a strategy to test how the brain accomplishes object-motion detection. Specifically, we tested the neural sensitivity of brain regions that are known to respond to egomotion-compatible visual motion (i.e., egomotion areas: cingulate sulcus visual area, posterior cingulate sulcus area, posterior insular cortex [PIC], V6+, V3A, IPSmot/VIP, and MT+) to a combination of different velocities of visually induced translational self- and object-motion within a virtual scene while participants were instructed to detect object-motion. To this aim, we combined individual surface-based brain mapping, task-evoked activity by functional magnetic resonance imaging, and parametric and representational similarity analyses. We found that all the egomotion regions (except area PIC) responded to all the possible combinations of self- and object-motion and were modulated by the self-motion velocity. Interestingly, we found that, among all the egomotion areas, only MT+, V6+, and V3A were further modulated by object-motion velocities, hence reflecting their possible role in discriminating between distinct velocities of self- and object-motion. We suggest that these egomotion regions may be involved in the complex computation required for detecting scene-relative object-motion during self-motion.

Effector-selective modulation of the effective connectivity within frontoparietal circuits during visuomotor tasks.

Despite extensive research, the functional architecture of the subregions of the dorsal posterior parietal cortex (PPC) involved in sensorimotor processing is far from clear. Here, we draw a thorough picture of the large-scale functional organization of the PPC to disentangle the fronto-parietal networks mediating visuomotor functions. To this aim, we reanalyzed available human functional magnetic resonance imaging data collected during the execution of saccades, hand, and foot pointing, and we combined individual surface-based activation, resting-state functional connectivity, and effective connectivity analyses. We described a functional distinction between a more lateral region in the posterior intraparietal sulcus (lpIPS), preferring saccades over pointing and coupled with the frontal eye fields (FEF) at rest, and a more medial portion (mpIPS) intrinsically correlated to the dorsal premotor cortex (PMd). Dynamic causal modeling revealed feedforward-feedback loops linking lpIPS with FEF during saccades and mpIPS with PMd during pointing, with substantial differences between hand and foot. Despite an intrinsic specialization of the action-specific fronto-parietal networks, our study reveals that their functioning is finely regulated according to the effector to be used, being the dynamic interactions within those networks differently modulated when carrying out a similar movement (i.e. pointing) but with distinct effectors (i.e. hand and foot).

Acute effects of neuromuscular electrical stimulation on cortical dynamics and reflex activation.

Superimposing neuromuscular electrical stimulation (NMES) on voluntary muscle contractions has shown the potential to improve motor performance even more than voluntary exercise alone. Nevertheless, the neurophysiological and neurocognitive mechanisms underlying this technique are still unclear. The aim of this study was to investigate the acute responses in spinal excitability and brain activity following three conditions: NMES superimposed on isometric contractions (NMES + ISO), passive NMES, and voluntary isometric contractions (ISO). Each condition involved 15 intermittent ankle plantar-flexions at submaximal level. Before and after each condition, tibial nerve stimulation was used to elicit H-reflexes, which represent a measure of spinal excitability, and somatosensory evoked potentials (SEPs), which index the activity of subcortical and cortical somatosensory areas. H-reflex amplitudes increased after NMES + ISO and decreased after passive NMES compared with baseline values, whereas they remained unaltered after ISO. Subcortical lemniscal activity remained unaltered after the three conditions. Activity in both primary and secondary somatosensory cortices (S1 and S2) increased after NMES + ISO and decreased after the ISO condition, whereas no differences emerged after NMES. At later stages of S2 processing, ISO induced no changes in cortical activity, which, conversely, increased after NMES and NMES + ISO. These findings indicate that the beneficial effects of NMES may be mediated by potentiation of the reflex pathways at the spinal level. At the brain level, peripheral input representation in the brain stem was not influenced by the experimental conditions, which, conversely, altered cortical activity by affecting synaptic efficiency through the somatosensory pathway.NEW & NOTEWORTHY Neuromuscular electrical stimulation superimposed on voluntary contractions (NMES+) is effective to improve motor performance in several populations. Here, we investigated the changes in cortical activation and reflex response following three acute conditions, including NMES+. Our results show that NMES+ has a greater excitatory effect at both spinal and cortical levels compared with passive stimulation and voluntary exercise alone. These results open up original perspectives for the implementation of NMES+ in neurorehabilitation and training environments.

Multisensory integration in cortical regions responding to locomotion-related visual and somatomotor signals.

During real-world locomotion, in order to be able to move along a path or avoid an obstacle, continuous changes in self-motion direction (i.e. heading) are needed. Control of heading changes during locomotion requires the integration of multiple signals (i.e., visual, somatomotor, vestibular). Recent fMRI studies have shown that both somatomotor areas (human PEc [hPEc], human PE [hPE], primary somatosensory cortex [S-I]) and egomotion visual regions (cingulate sulcus visual area [CSv], posterior cingulate area [pCi], posterior insular cortex [PIC]) respond to either leg movements and egomotion-compatible visual stimulations, suggesting a role in the analysis of both visual attributes of egomotion and somatomotor signals with the aim of guiding locomotion. However, whether these regions are able to integrate egomotion-related visual signals with somatomotor inputs coming from leg movements during heading changes remains an open question. Here we used a combined approach of individual functional localizers and task-evoked activity by fMRI. In thirty subjects we first localized three egomotion areas (CSv, pCi, PIC) and three somatomotor regions (S-I, hPE, hPEc). Then, we tested their responses in a multisensory integration experiment combining visual and somatomotor signals relevant to locomotion in congruent or incongruent trials. We used an fMR-adaptation paradigm to explore the sensitivity to the repeated presentation of these bimodal stimuli in the six regions of interest. Results revealed that hPE, S-I and CSv showed an adaptation effect regardless of congruency, while PIC, pCi and hPEc showed sensitivity to congruency. PIC exhibited a preference for congruent trials compared to incongruent trials. Areas pCi and hPEc exhibited an adaptation effect only for congruent and incongruent trials, respectively. PIC, pCi and hPEc sensitivity to the congruency relationship between visual (locomotion-compatible) cues and (leg-related) somatomotor inputs suggests that these regions are involved in multisensory integration processes, likely in order to guide/adjust leg movements during heading changes.

Effect of Exoskeleton-Assisted Rehabilitation Over Prefrontal Cortex in Multiple Sclerosis Patients: A Neuroimaging Pilot Study.

Application of a passive and fully articulated exoskeleton, called Human Body Posturizer (HBP), has been demonstrated to improve mobility, response accuracy and ambulation in multiple sclerosis (MS) patients. By using functional magnetic imaging (fMRI) during a visuomotor discrimination task, we performed a pilot study to evaluate the effect of HBP over the neural correlates of motor and cognitive functions which are typically impaired in MS patients. Specifically, we tested the effect of a 6-week multidisciplinary rehabilitation intervention on two groups of MS patients: a control group who followed a standard physiotherapeutic rehabilitation protocol, and an experimental group who used the HBP during physical exercises in addition to the standard protocol. We found that, after treatment, the experimental group exhibited a significant lower activity (as compared to the control group) in the inferior frontal gyrus. This post-treatment activity reduction can be explained as a retour to a normal range, being the amount of iFg activity observed in the experimental patients very similar to that observed in healthy subjects. These findings indicate that the use of HBP during rehabilitation intervention normalizes the prefrontal activity, mitigating the cortical hyperactivity associated to MS.

Neural bases of self- and object-motion in a naturalistic vision.

To plan movements toward objects our brain must recognize whether retinal displacement is due to self-motion and/or to object-motion. Here, we aimed to test whether motion areas are able to segregate these types of motion. We combined an event-related functional magnetic resonance imaging experiment, brain mapping techniques, and wide-field stimulation to study the responsivity of motion-sensitive areas to pure and combined self- and object-motion conditions during virtual movies of a train running within a realistic landscape. We observed a selective response in MT to the pure object-motion condition, and in medial (PEc, pCi, CSv, and CMA) and lateral (PIC and LOR) areas to the pure self-motion condition. Some other regions (like V6) responded more to complex visual stimulation where both object- and self-motion were present. Among all, we found that some motion regions (V3A, LOR, MT, V6, and IPSmot) could extract object-motion information from the overall motion, recognizing the real movement of the train even when the images remain still (on the screen), or moved, because of self-movements. We propose that these motion areas might be good candidates for the "flow parsing mechanism," that is the capability to extract object-motion information from retinal motion signals by subtracting out the optic flow components.

Prompting future events: Effects of temporal cueing and time on task on brain preparation to action.

Prediction about event timing plays a leading role in organizing and optimizing behavior. We recorded anticipatory brain activities and evaluated whether temporal orienting processes are reflected by the novel prefrontal negative (pN) component, as already shown for the contingent negative variation (CNV). Fourteen young healthy participants underwent EEG and fMRI recordings in separate sessions; they were asked to perform a Go/No-Go task in which temporal orienting was manipulated: the external condition (a visual display indicating the time of stimulus onset) and the internal condition (time information not provided). In both conditions, the source of the pN was localized in the pars opercularis of the iFg; the source of the CNV was localized in the supplementary motor area and cingulate motor area, as expected. Anticipatory activity was also found in the occipital-parietal cortex. Time on task EEG analysis showed a marked learning effect in the internal condition, while the effect was minor in the external condition. In fMRI, the two conditions had a similar pattern; similarities and differences of results obtained with the two techniques are discussed. Overall, data are consistent with the view that the pN reflects a proactive cognitive control, including temporal orienting.

A putative human homologue of the macaque area PEc.

The cortical area PEc is anatomically and functionally well-defined in macaque, but it is unknown whether it has a counterpart in human. Since we know that macaque PEc, but not the nearby posterior regions, hosts a lower limb representation, in an attempt to recognize a possible human PEc we looked for the existence of leg representations in the human parietal cortex using individual cortical surface-based analysis, task-evoked paradigms and resting-state functional connectivity. fMRI images were acquired while thirty-one participants performed long-range leg movements through an in-house MRI-compatible set-up. We revealed the existence of multiple leg representations in the human dorsomedial parietal cortex, here defined as S-I (somatosensory-I), hPE (human PE, in the postcentral sulcus), and hPEc (human PEc, in the anterior precuneus). Among the three "leg" regions, hPEc had a unique functional profile, in that it was the only one responding to both arm and leg movements, to both hand-pointing and foot pointing movements, and to flow field visual stimulation, very similar to macaque area PEc. In addition, hPEc showed functional connections with the somatomotor regions hosting a lower limb representation, again as in macaque area PEc. Therefore, based on similarity in brain position, functional organization, cortical connections, and relationship with the neighboring areas, we propose that this cortical region is the human homologue of macaque area PEc.

Egomotion-related visual areas respond to active leg movements.

Monkey neurophysiology and human neuroimaging studies have demonstrated that passive viewing of optic flow stimuli activates a cortical network of temporal, parietal, insular, and cingulate visual motion regions. Here, we tested whether the human visual motion areas involved in processing optic flow signals simulating self-motion are also activated by active lower limb movements, and hence are likely involved in guiding human locomotion. To this aim, we used a combined approach of task-evoked activity and resting-state functional connectivity by fMRI. We localized a set of six egomotion-responsive visual areas (V6+, V3A, intraparietal motion/ventral intraparietal [IPSmot/VIP], cingulate sulcus visual area [CSv], posterior cingulate sulcus area [pCi], posterior insular cortex [PIC]) by using optic flow. We tested their response to a motor task implying long-range active leg movements. Results revealed that, among these visually defined areas, CSv, pCi, and PIC responded to leg movements (visuomotor areas), while V6+, V3A, and IPSmot/VIP did not (visual areas). Functional connectivity analysis showed that visuomotor areas are connected to the cingulate motor areas, the supplementary motor area, and notably to the medial portion of the somatosensory cortex, which represents legs and feet. We suggest that CSv, pCi, and PIC perform the visual analysis of egomotion-like signals to provide sensory information to the motor system with the aim of guiding locomotion.

Immediate cortical adaptation in visual and non-visual areas functions induced by monovision.

KEY POINTS: Monovision is an optical correction for presbyopes that consists of correcting one eye for far distance and the other for near distance, creating a superimposition of an in-focus with a blurred image. Brain adaptation to monovision was studied in unexperienced observers by measuring visual evoked potentials from 64-channels. The first clear effect of monovision on visual evoked potentials was the C1 amplitude reduction, indicating that the unilateral blurring induced by monovision reduces feed-forward activity in primary visual area. Monovision led also to an increased amplitude of the P1 and pP1 components, with the latter originating in prefrontal regions. This effect probably works as an attentional compensatory activity used to compensate for the degraded V1 signal. ABSTRACT: A common and often successful option to correct presbyopia with contact lenses is monovision. This is an unbalanced correction across the two eyes where one eye is corrected for far vision and the other eye is corrected for near vision. Monovision is therefore a form of acquired anisometropia that causes a superimposition of an in-focus image with a blurred image. In spite of this visual anisometropia, monovision has been successfully used for many decadesl however the brain mechanism supporting monovision is not well understood. The present study aimed to measure the visual evoked potentials with a high-density electrode array (64-channel) in a group of presbyopes and to provide a detailed spatiotemporal analysis of the cortical activity after a short period of adaptation to monovision with contact lenses. When compared with a balanced eye near correction, monovision produced both a clear reduction of the earliest visual evoked potential components, the C1 and the N1, and an amplitude increase of the P1 and pP1. These results indicate that the unilateral blurring induced by wearing monovision contact lenses reduces feed-forward activity in the primary visual area and feedback activity in extrastriate areas (C1 and N1 reduction). Interestingly, other brain activities in both extrastriate visual areas (the P1 component) and in the anterior insula (the pP1 component) appear to compensate for this dysfunction, increasing their activity during monovision. These changes confirm the presence of fluid brain adaptation in visual and non-visual areas during monocular interferences.

Detailed spatiotemporal brain mapping of chromatic vision combining high-resolution VEP with fMRI and retinotopy.

Neuroimaging studies have identified so far, several color-sensitive visual areas in the human brain, and the temporal dynamics of these activities have been separately investigated using the visual-evoked potentials (VEPs). In the present study, we combined electrophysiological and neuroimaging methods to determine a detailed spatiotemporal profile of chromatic VEP and to localize its neural generators. The accuracy of the present co-registration study was obtained by combining standard fMRI data with retinotopic and motion mapping data at the individual level. We found a sequence of occipito activities more complex than that typically reported for chromatic VEPs, including feed-forward and reentrant feedback. Results showed that chromatic human perception arises by the combined activity of at the least five parieto-occipital areas including V1, LOC, V8/VO, and the motion-sensitive dorsal region MT+. However, the contribution of V1 and V8/VO seems dominant because the re-entrant activity in these areas was present more than once (twice in V8/VO and thrice in V1). This feedforward and feedback chromatic processing appears delayed compared with the luminance processing. Associating VEPs and neuroimaging measures, we showed for the first time a complex spatiotemporal pattern of activity, confirming that chromatic stimuli produce intricate interactions of many different brain dorsal and ventral areas.

Hemispheric asymmetries in the transition from action preparation to execution.

Flexible and adaptive behavior requires the ability to contextually stop inappropriate actions and select the right one as quickly as possible. Recently, it has been proposed that three brain regions, i.e., the inferior frontal gyrus (iFg), the anterior insula (aIns), and the anterior intraparietal sulcus (aIPs), play an important role in several processing phases of perceptual decision tasks, especially in the preparation, perception and action phases, respectively. However, little is known about hemispheric differences in the activation of these three areas during the transition from perception to action. Many studies have examined how people prepare to stop upcoming responses through both proactive and reactive inhibitory control. Although inhibitory control has been associated with activity in the right prefrontal cortex (PFC), we have previously reported that, during a discriminative response task performed with the right hand, we observed: 1) a bilateral activity in the iFg during the preparation phase, and 2) a left dominant activity in the aIns and aIPs during the transition from perception to action, i.e., the so-called stimulus-response mapping. To clarify the hemispheric dominance of these processes, we combined the high temporal resolution of event-related potentials (ERPs) with the high spatial resolution of event-related functional magnetic resonance imaging (fMRI) while participants performed a discriminative response task (DRT) and a simple response task (SRT) using their non-dominant left hand. We confirmed that proactive inhibitory control originates in the iFg: its activity started one second before the stimulus onset and was released concomitantly to the stimulus appearance. Most importantly, we confirmed the presence of a bilateral iFg activity that seems to reflect a bilateral proactive control rather than a right-hemisphere dominance or a stronger control of the hemisphere contralateral to the responding hand. Further, we observed a stronger activation of the left aIns and a right-lateralized activation of the aIPs reflecting left-hemisphere dominance for stimulus-response mapping finalized to response execution and a contralateral-hand parietal premotor activity, respectively.

Adaptive smoothing based on Gaussian processes regression increases the sensitivity and specificity of fMRI data.

Temporal and spatial filtering of fMRI data is often used to improve statistical power. However, conventional methods, such as smoothing with fixed-width Gaussian filters, remove fine-scale structure in the data, necessitating a tradeoff between sensitivity and specificity. Specifically, smoothing may increase sensitivity (reduce noise and increase statistical power) but at the cost loss of specificity in that fine-scale structure in neural activity patterns is lost. Here, we propose an alternative smoothing method based on Gaussian processes (GP) regression for single subjects fMRI experiments. This method adapts the level of smoothing on a voxel by voxel basis according to the characteristics of the local neural activity patterns. GP-based fMRI analysis has been heretofore impractical owing to computational demands. Here, we demonstrate a new implementation of GP that makes it possible to handle the massive data dimensionality of the typical fMRI experiment. We demonstrate how GP can be used as a drop-in replacement to conventional preprocessing steps for temporal and spatial smoothing in a standard fMRI pipeline. We present simulated and experimental results that show the increased sensitivity and specificity compared to conventional smoothing strategies. Hum Brain Mapp 38:1438-1459, 2017. © 2016 Wiley Periodicals, Inc.

Spatiotemporal brain mapping during preparation, perception, and action.

Deciding whether to act or not to act is a fundamental cognitive function. To avoid incorrect responses, both reactive and proactive modes of control have been postulated. Little is known, however, regarding the brain implementation of proactive mechanisms, which are deployed prior to an actual need to inhibit a response. Via a combination of electrophysiological and neuroimaging measures (recorded in 21 and 16 participants, respectively), we describe the brain localization and timing of neural activity that underlies the anticipatory proactive mechanism. From these results, we conclude that proactive control originates in the inferior Frontal gyrus, is established well before stimulus perception, and is released concomitantly with stimulus appearance. Stimulus perception triggers early activity in the anterior insula and intraparietal cortex contralateral to the responding hand; these areas likely mediate the transition from perception to action. The neural activities leading to the decision to act or not to act are described in the framework of a three-stage model that includes perception, action, and anticipatory functions taking place well before stimulus onset.

The human cortical areas V6 and V6A.

In macaque, it has long been known since the late nineties that the medial parieto-occipital sulcus (POS) contains two regions, V6 and V6A, important for visual motion and action. While V6 is a retinotopically organized extrastriate area, V6A is a broadly retinotopically organized visuomotor area constituted by a ventral and dorsal subdivision (V6Av and V6Ad), both containing arm movement-related cells active during spatially directed reaching movements. In humans, these areas have been mapped only in recent years thanks to neuroimaging methods. In a series of brain mapping studies, by using a combination of functional magnetic resonance imaging methods such as wide-field retinotopy and task-evoked activity, we mapped human areas V6 (Pitzalis et al., 2006) and V6Av (Pitzalis et al., 2013 d) retinotopically and defined human V6Ad functionally as a pointing-selective region situated anteriorly in the close proximity of V6Av (Tosoni et al., 2014). Like in macaque, human V6 is a motion area (e.g., Pitzalis et al., 2010, 2012, 2013 a, b , c ), while V6Av and V6Ad respond to pointing movements (Tosoni et al., 2014). The retinotopic organization (when present), anatomical position, neighbor relations, and functional properties of these three areas closely resemble those reported for macaque V6 (Galletti et al., 1996, 1999 a), V6Av, and V6Ad (Galletti et al., 1999 b; Gamberini et al., 2011). We suggest that information on objects in depth which are translating in space, because of the self-motion, is processed in V6 and conveyed to V6A for evaluating object distance in a dynamic condition such as that created by self-motion, so to orchestrate the eye and arm movements necessary to reach or avoid static and moving objects in the environment.

Common and specific activations supporting optic flow processing and navigation as revealed by a meta-analysis of neuroimaging studies.

Optic flow provides useful information in service of spatial navigation. However, whether brain networks supporting these two functions overlap is still unclear. Here we used Activation Likelihood Estimation (ALE) to assess the correspondence between brain correlates of optic flow processing and spatial navigation and their specific neural activations. Since computational and connectivity evidence suggests that visual input from optic flow provides information mainly during egocentric navigation, we further tested the correspondence between brain correlates of optic flow processing and that of both egocentric and allocentric navigation. Optic flow processing shared activation with egocentric (but not allocentric) navigation in the anterior precuneus, suggesting its role in providing information about self-motion, as derived from the analysis of optic flow, in service of egocentric navigation. We further documented that optic flow perception and navigation are partially segregated into two functional and anatomical networks, i.e., the dorsal and the ventromedial networks. Present results point to a dynamic interplay between the dorsal and ventral visual pathways aimed at coordinating visually guided navigation in the environment.

Cortical dynamics in visual areas induced by the first use of multifocal contact lenses in presbyopes.

A common non-spectacle strategy to correct presbyopia is to provide simultaneous images with multifocal optical designs. Understanding the neuroadaptation mechanisms behind multifocal devices usage would have important clinical implications, such as predicting whether patients will be able to tolerate multifocal optics. The aim of this study was to evaluate the brain correlates during the initial wear of multifocal contact lenses (CLs) using high-density visual evoked potential (VEP) measures. Fifteen presbyopes (mean age 51.8 ±â€¯2.6 years) who had previously not used multifocal CLs were enrolled. VEP measures were achieved while participants looked at arrays of 0.5 logMAR Sloan letters in three different optical conditions arranged with CLs: monofocal condition with the optical power appropriate for the distance viewing; multifocal correction with medium addition; and multifocal correction with low addition. An ANOVA for repeated measures showed that the amplitude of the C1 and N1 components significantly dropped with both multifocal low and medium addition CL conditions compared to monofocal CLs. The P1 and P2 components showed opposite behavior with an increase in amplitudes for multifocal compared to monofocal conditions. VEP data indicated that multifocal presbyopia corrections produce a loss of feedforward activity in the primary visual cortex that is compensated by extra feedback activity in extrastriate areas only, in both early and late visual processing.

Parallel motion signals to the medial and lateral motion areas V6 and MT+.

MT+ and V6 are key motion areas of the dorsal visual stream in both macaque and human brains. In the present study, we combined electrophysiological and neuroimaging methods (including retinotopic brain mapping) to find the electrophysiological correlates of V6 and to define its temporal relationship with the activity observed in MT+. We also determined the spatio-temporal profile of the motion coherency effect on visual evoked potentials (VEPs), and localized its neural generators. We found that area V6 participates in the very early phase of the coherent motion processing and that its electroencephalographic activity is almost simultaneous with that of MT+. We also found a late second activity in V6 that we interpret as a re-entrant feedback from extrastriate visual areas (e.g. area V3A). Three main cortical sources were differently modulated by the motion coherence: while V6 and MT+ showed a preference for the coherent motion, area V3A preferred the random condition. The response timing of these cortical sources indicates that motion signals flow in parallel from the occipital pole to the medial and lateral motion areas V6 and MT+, suggesting the view of a differential functional role.

Functional organization of the caudal part of the human superior parietal lobule.

Like in macaque, the caudal portion of the human superior parietal lobule (SPL) plays a key role in a series of perceptive, visuomotor and somatosensory processes. Here, we review the functional properties of three separate portions of the caudal SPL, i.e., the posterior parieto-occipital sulcus (POs), the anterior POs, and the anterior part of the caudal SPL. We propose that the posterior POs is mainly dedicated to the analysis of visual motion cues useful for object motion detection during self-motion and for spatial navigation, while the more anterior parts are implicated in visuomotor control of limb actions. The anterior POs is mainly involved in using the spotlight of attention to guide reach-to-grasp hand movements, especially in dynamic environments. The anterior part of the caudal SPL plays a central role in visually guided locomotion, being implicated in controlling leg-related movements as well as the four limbs interaction with the environment, and in encoding egomotion-compatible optic flow. Together, these functions reveal how the caudal SPL is strongly implicated in skilled visually-guided behaviors.

Spatiotemporal brain mapping of spatial attention effects on pattern-reversal ERPs.

Recordings of event-related potentials (ERPs) were combined with structural and functional magnetic resonance imaging (fMRI) to investigate the timing and localization of stimulus selection processes during visual-spatial attention to pattern-reversing gratings. Pattern reversals were presented in random order to the left and right visual fields at a rapid rate, while subjects attended to the reversals in one field at a time. On separate runs, stimuli were presented in the upper and lower visual quadrants. The earliest ERP component (C1, peaking at around 80 ms), which inverted in polarity for upper versus lower field stimuli and was localized in or near visual area V1, was not modulated by attention. In the latency range 80-250 ms, multiple components were elicited that were increased in amplitude by attention and were colocalized with fMRI activations in specific visual cortical areas. The principal anatomical sources of these attention-sensitive components were localized by fMRI-seeded dipole modeling as follows: P1 (ca. 100 ms-source in motion-sensitive area MT+), C2 (ca. 130 ms-same source as C1), N1a (ca. 145 ms-source in horizontal intraparietal sulcus), N1b (ca. 165 ms-source in fusiform gyrus, area V4/V8), N1c (ca. 180 ms-source in posterior intraparietal sulcus, area V3A), and P2 (ca. 220 ms-multiple sources, including parieto-occipital sulcus, area V6). These results support the hypothesis that spatial attention acts to amplify both feed-forward and feedback signals in multiple visual areas of both the dorsal and ventral streams of processing.