Perceptual-Cognitive Integration for Goal-Directed Action in Naturalistic Environments.

Real-world actions require one to simultaneously perceive, think, and act on the surrounding world, requiring the integration of (bottom-up) sensory information and (top-down) cognitive and motor signals. Studying these processes involves the intellectual challenge of cutting across traditional neuroscience silos, and the technical challenge of recording data in uncontrolled natural environments. However, recent advances in techniques, such as neuroimaging, virtual reality, and motion tracking, allow one to address these issues in naturalistic environments for both healthy participants and clinical populations. In this review, we survey six topics in which naturalistic approaches have advanced both our fundamental understanding of brain function and how neurologic deficits influence goal-directed, coordinated action in naturalistic environments. The first part conveys fundamental neuroscience mechanisms related to visuospatial coding for action, adaptive eye-hand coordination, and visuomotor integration for manual interception. The second part discusses applications of such knowledge to neurologic deficits, specifically, steering in the presence of cortical blindness, impact of stroke on visual-proprioceptive integration, and impact of visual search and working memory deficits. This translational approach-extending knowledge from lab to rehab-provides new insights into the complex interplay between perceptual, motor, and cognitive control in naturalistic tasks that are relevant for both basic and clinical research.

Influence of Gaze, Vision, and Memory on Hand Kinematics in a Placement Task.

People usually reach for objects to place them in some position and orientation, but the placement component of this sequence is often ignored. For example, reaches are influenced by gaze position, visual feedback, and memory delays, but their influence on object placement is unclear. Here, we tested these factors in a task where participants placed and oriented a trapezoidal block against 2D visual templates displayed on a frontally located computer screen. In Experiment 1, participants matched the block to three possible orientations: 0o (horizontal), +45o and -45o, with gaze fixated 10o to the left/right. The hand and template either remained illuminated (closed-loop), or visual feedback was removed (open-loop). Here, hand location consistently overshot the template relative to gaze, especially in the open-loop task; likewise, orientation was influenced by gaze position (depending on template orientation and visual feedback). In Experiment 2, a memory delay was added, and participants sometimes performed saccades (towards, away from, or across the template). In this task, the influence of gaze on orientation vanished, but location errors were influenced by both template orientation and final gaze position. Contrary to our expectations, the previous saccade metrics also impacted placement overshoot. Overall, hand orientation was influenced by template orientation in a nonlinear fashion. These results demonstrate interactions between gaze and orientation signals in the planning and execution of hand placement and suggest different neural mechanisms for closed-loop, open-loop, and memory delay placement.

Angular Correlations in the ß Decay of

We present the first measurement of the α-ß-ν angular correlation in the Gamow-Teller ß^{+} decay of ^{8}B. This was accomplished using the Beta-decay Paul Trap, expanding on our previous work on the ß^{-} decay of ^{8}Li. The ^{8}B result is consistent with the V-A electroweak interaction of the standard model and, on its own, provides a limit on the exotic right-handed tensor current relative to the axial-vector current of |C_{T}/C_{A}|^{2}<0.013 at the 95.5% confidence level. This represents the first high-precision angular correlation measurements in mirror decays and was made possible through the use of an ion trap. By combining this ^{8}B result with our previous ^{8}Li results, we demonstrate a new pathway for increased precision in searches for exotic currents.

Parietofrontal oscillations show hand-specific interactions with top-down movement plans.

To generate a hand-specific reach plan, the brain must integrate hand-specific signals with the desired movement strategy. Although various neurophysiology/imaging studies have investigated hand-target interactions in simple reach-to-target tasks, the whole brain timing and distribution of this process remain unclear, especially for more complex, instruction-dependent motor strategies. Previously, we showed that a pro/anti pointing instruction influences magnetoencephalographic (MEG) signals in frontal cortex that then propagate recurrently through parietal cortex (Blohm G, Alikhanian H, Gaetz W, Goltz HC, DeSouza JF, Cheyne DO, Crawford JD. NeuroImage 197: 306-319, 2019). Here, we contrasted left versus right hand pointing in the same task to investigate 1) which cortical regions of interest show hand specificity and 2) which of those areas interact with the instructed motor plan. Eight bilateral areas, the parietooccipital junction (POJ), superior parietooccipital cortex (SPOC), supramarginal gyrus (SMG), medial/anterior interparietal sulcus (mIPS/aIPS), primary somatosensory/motor cortex (S1/M1), and dorsal premotor cortex (PMd), showed hand-specific changes in beta band power, with four of these (M1, S1, SMG, aIPS) showing robust activation before movement onset. M1, SMG, SPOC, and aIPS showed significant interactions between contralateral hand specificity and the instructed motor plan but not with bottom-up target signals. Separate hand/motor signals emerged relatively early and lasted through execution, whereas hand-motor interactions only occurred close to movement onset. Taken together with our previous results, these findings show that instruction-dependent motor plans emerge in frontal cortex and interact recurrently with hand-specific parietofrontal signals before movement onset to produce hand-specific motor behaviors.NEW & NOTEWORTHY The brain must generate different motor signals depending on which hand is used. The distribution and timing of hand use/instructed motor plan integration are not understood at the whole brain level. Using MEG we show that different action planning subnetworks code for hand usage and integrating hand use into a hand-specific motor plan. The timing indicates that frontal cortex first creates a general motor plan and then integrates hand specificity to produce a hand-specific motor plan.

Action planning modulates the representation of object features in human fronto-parietal and occipital cortex.

The visual cortex has been extensively studied to investigate its role in object recognition but to a lesser degree to determine how action planning influences the representation of objects' features. We used functional MRI and pattern classification methods to determine if during action planning, object features (orientation and location) could be decoded in an action-dependent way. Sixteen human participants used their right dominant hand to perform movements (Align or Open reach) towards one of two 3D-real oriented objects that were simultaneously presented and placed on either side of a fixation cross. While both movements required aiming towards target location, Align but not Open reach movements required participants to precisely adjust hand orientation. Therefore, we hypothesized that if the representation of object features is modulated by the upcoming action, pre-movement activity pattern would allow more accurate dissociation between object features in Align than Open reach tasks. We found such dissociation in the anterior and posterior parietal cortex, as well as in the dorsal premotor cortex, suggesting that visuomotor processing is modulated by the upcoming task. The early visual cortex showed significant decoding accuracy for the dissociation between object features in the Align but not Open reach task. However, there was no significant difference between the decoding accuracy in the two tasks. These results demonstrate that movement-specific preparatory signals modulate object representation in the frontal and parietal cortex, and to a lesser extent in the early visual cortex, likely through feedback functional connections.

Iris and Ciliary Body Melanocytomas Are Defined by Solitary GNAQ Mutation Without Additional Oncogenic Alterations.

OBJECTIVE: To analyze the genetic features of melanocytomas and melanomas of the anterior uvea and assess the value of molecular testing for diagnosis and prognostication. DESIGN: Retrospective case-control study. SUBJECTS: Patients with melanocytoma (n = 16) and melanoma (n = 19) of the anterior uvea. METHODS: Targeted next-generation sequencing was performed on formalin-fixed, paraffin-embedded tumor tissue from anterior uveal melanocytic tumors and correlated with clinicopathologic features. MAIN OUTCOME MEASURES: Presence or absence of accompanying oncogenic alterations beyond GNAQ/GNA11 and their association with histologic features and local recurrence. RESULTS: Hotspot missense mutations in GNAQ/GNA11 were identified in 91% (32/35) of all cases. None of the melanocytomas with or without atypia demonstrated chromosomal imbalances or additional oncogenic variants beyond GNAQ mutation, and none recurred over a median follow-up of 36 months. Additional alterations identified in a subset of melanomas include mutations in BAP1 (n = 3), EIF1AX (n = 4), SRSF2 (n = 1), PTEN (n = 1), and EP300 (n = 1); monosomy 3p (n = 6); trisomy 6p (n = 3); trisomy 8q (n = 2); and an ultraviolet mutational signature (n = 5). Local recurrences were limited to melanomas, all of which demonstrated oncogenic alterations in addition to GNAQ/GNA11 (n = 5). A single melanoma harboring GNAQ and BAP1 mutations and monosomy 3 was the only tumor that metastasized. CONCLUSIONS: In this study, anterior segment uveal melanocytomas did not display oncogenic alterations beyond GNAQ/GNA11. Therefore, they are genetically similar to uveal nevi rather than uveal melanoma based on their molecular features known from the literature. Molecular testing can be performed on borderline cases to aid risk stratification and clinical management decisions.

Parietal Cortex Integrates Saccade and Object Orientation Signals to Update Grasp Plans.

Coordinated reach-to-grasp movements are often accompanied by rapid eye movements (saccades) that displace the desired object image relative to the retina. Parietal cortex compensates for this by updating reach goals relative to current gaze direction, but its role in the integration of oculomotor and visual orientation signals for updating grasp plans is unknown. Based on a recent perceptual experiment, we hypothesized that inferior parietal cortex (specifically supramarginal gyrus [SMG]) integrates saccade and visual signals to update grasp plans in additional intraparietal/superior parietal regions. To test this hypothesis in humans (7 females, 6 males), we used a functional magnetic resonance paradigm, where saccades sometimes interrupted grasp preparation toward a briefly presented object that later reappeared (with the same/different orientation) just before movement. Right SMG and several parietal grasp regions, namely, left anterior intraparietal sulcus and bilateral superior parietal lobule, met our criteria for transsaccadic orientation integration: they showed task-dependent saccade modulations and, during grasp execution, they were specifically sensitive to changes in object orientation that followed saccades. Finally, SMG showed enhanced functional connectivity with both prefrontal saccade regions (consistent with oculomotor input) and anterior intraparietal sulcus/superior parietal lobule (consistent with sensorimotor output). These results support the general role of parietal cortex for the integration of visuospatial perturbations, and provide specific cortical modules for the integration of oculomotor and visual signals for grasp updating.SIGNIFICANCE STATEMENT How does the brain simultaneously compensate for both external and internally driven changes in visual input? For example, how do we grasp an unstable object while eye movements are simultaneously changing its retinal location? Here, we used fMRI to identify a group of inferior parietal (supramarginal gyrus) and superior parietal (intraparietal and superior parietal) regions that show saccade-specific modulations during unexpected changes in object/grasp orientation, and functional connectivity with frontal cortex saccade centers. This provides a network, complementary to the reach goal updater, that integrates visuospatial updating into grasp plans, and may help to explain some of the more complex symptoms associated with parietal damage, such as constructional ataxia.

Neuromagnetic signatures of the spatiotemporal transformation for manual pointing.

Movement planning involves transforming the sensory signals into a command in motor coordinates. Surprisingly, the real-time dynamics of sensorimotor transformations at the whole brain level remain unknown, in part due to the spatiotemporal limitations of fMRI and neurophysiological recordings. Here, we used magnetoencephalography (MEG) during pro-/anti-wrist pointing to determine (1) the cortical areas involved in transforming visual signals into appropriate hand motor commands, and (2) how this transformation occurs in real time, both within and across the regions involved. We computed sensory, motor, and sensorimotor indices in 16 bilateral brain regions for direction coding based on hemispherically lateralized de/synchronization in the α (7-15 Hz) and ß (15-35 Hz) bands. We found a visuomotor progression, from pure sensory codes in 'early' occipital-parietal areas, to a temporal transition from sensory to motor coding in the majority of parietal-frontal sensorimotor areas, to a pure motor code, in both the α and ß bands. Further, the timing of these transformations revealed a top-down pro/anti cue influence that propagated 'backwards' from frontal through posterior cortical areas. These data directly demonstrate a progressive, real-time transformation both within and across the entire occipital-parietal-frontal network that follows specific rules of spatial distribution and temporal order.

The role of pallidum in the neural integrator model of cervical dystonia.

Dystonia is the third most common movement disorder affecting three million people worldwide. Cervical dystonia is the most common form of dystonia. Despite common prevalence the pathophysiology of cervical dystonia is unclear. Traditional view is that basal ganglia is involved in pathophysiology of cervical dystonia, while contemporary theories suggested the role of cerebellum and proprioception in the pathophysiology of cervical dystonia. It was recently proposed that the cervical dystonia is due to malfunctioning of the head neural integrator - the neuron network that normally converts head velocity to position. Most importantly the neural integrator model was inclusive of traditional proposal emphasizing the role of basal ganglia while also accommodating the contemporary view suggesting the involvement of cerebellum and proprioception. It was hypothesized that the head neural integrator malfunction is the result of impairment in cerebellar, basal ganglia, or proprioceptive feedback that converge onto the integrator. The concept of converging input from the basal ganglia, cerebellum, and proprioception to the network participating in head neural integrator explains that abnormality originating anywhere in the network can lead to the identical motor deficits - drifts followed by rapid corrective movements - a signature of neural integrator dysfunction. We tested this hypothesis in an experiment examining simultaneously recorded globus pallidal single-unit activity, synchronized neural activity (local field potential), and electromyography (EMG) measured from the neck muscles during the standard-of-care deep brain stimulation surgery in 12 cervical dystonia patients (24 hemispheres). Physiological data were collected spontaneously or during voluntary shoulder shrug activating the contralateral trapezius muscle. The activity of pallidal neurons during shoulder shrug exponentially decayed with time constants that were comparable to one measured from the pretectal neural integrator and the trapezius electromyography. These results show that evidence of abnormal neural integration is also seen in globus pallidum, and that latter is connected with the neural integrator. Pretectal single neuron responses consistently preceded the muscle activity; while the globus pallidum internus response always lagged behind the muscle activity. Globus pallidum externa had equal proportion of lag and lead neurons. These results suggest globus pallidum receive feedback from the muscles or the efference copy from the integrator or the other source of the feedback. There was bi-hemispheric asymmetry in the pallidal single-unit activity and local field potentials. The asymmetry correlated with degree of lateral head turning in cervical dystonia patients. These results suggest that bihemispheric asymmetry in the feedback leads to asymmetric dysfunction in the neural integrator causing head turning.

Three-dimensional transformations for goal-directed action.

Much of the central nervous system is involved in visuomotor transformations for goal-directed gaze and reach movements. These transformations are often described in terms of stimulus location, gaze fixation, and reach endpoints, as viewed through the lens of translational geometry. Here, we argue that the intrinsic (primarily rotational) 3-D geometry of the eye-head-reach systems determines the spatial relationship between extrinsic goals and effector commands, and therefore the required transformations. This approach provides a common theoretical framework for understanding both gaze and reach control. Combined with an assessment of the behavioral, neurophysiological, imaging, and neuropsychological literature, this framework leads us to conclude that (a) the internal representation and updating of visual goals are dominated by gaze-centered mechanisms, but (b) these representations must then be transformed as a function of eye and head orientation signals into effector-specific 3-D movement commands.

Multicenter Clinical Evaluation of the Automated Aries Group A Strep PCR Assay from Throat Swabs.

Group A Streptococcus (GAS) is one of the leading causes of bacterial pharyngitis. Early GAS diagnosis is critical for appropriate antibiotic administration that reduces the risk of GAS sequelae and limits spread of the infection. The Aries Group A Strep (GAS) assay (Luminex, Austin, TX) is a fully automated PCR assay for direct detection of GAS in throat swab specimens in less than 2 h with minimum hands-on time. This multicenter prospective study evaluated the clinical performance of the Aries GAS assay compared to that of Streptococcus pyogenes culture. Subjects with symptoms consistent with pharyngitis were enrolled across four sites in the United States, and a throat swab in liquid Amies medium was obtained. Aries and reference testing was performed within 72 and 48 h after sample collection, respectively. Of 623 throat swab specimens from patients with pharyngitis (93.6% <18 years old, 54.3% female), the reference method yielded valid results for 618 specimens. Reference and Aries assay testing showed GAS-positive results for 160 (25.9%) and 166 (26.9%) specimens, respectively. Compared to the reference method, Aries assay sensitivity was 97.5% (95% confidence interval [CI], 93.7% to 99.0%), specificity was 97.8% (95% CI, 96.0 to 98.8%), positive predictive value was 94.0% (95% CI, 89.3% to 96.7%), and negative predictive value was 99.1% (95% CI, 97.7% to 99.7%). There were 10 false-positive and four false-negative detections with the Aries assay. Discrepant analysis with bidirectional sequencing yielded concordant results with the Aries assay for nine of 14 discordant samples. The Aries assay had high sensitivity and specificity for qualitative detection of group A Streptococcus from patients with pharyngitis.

The Vascular Behavioral and Cognitive Disorders criteria for vascular cognitive disorders: a validation study.

BACKGROUND AND PURPOSE: The Vascular Behavioral and Cognitive Disorders (VASCOG) criteria for vascular cognitive disorders were published in 2014, but their concurrent and predictive validity have not been examined. METHODS: Participants (N = 165, aged 49-86 years) were from Sydney Stroke Study, a longitudinal study of post-stroke cognitive impairment and dementia. Diagnoses using the National Institute of Neurological Disorders and Stroke - Association Internationale pour la Recherché et l'Enseignement en Neurosciences (NINDS-AIREN), the Alzheimer's Disease Diagnostic and Treatment Centers (ADDTC) and the Diagnostic and Statistical Manual of Mental Disorders, 4th edition (DSM-IV), criteria for vascular dementia (VaD) were made by consensus at multidisciplinary case conferences. Diagnoses for mild vascular cognitive disorder (mVCD) and VaD using VASCOG, DSM-5 and the Vascular Impairment of Cognition Classification Consensus Study (VICCCS) criteria were made by two study authors. Agreement levels between criteria sets were examined using Cohen's kappa (κ). The ability of VaD diagnoses to predict mortality over 10 years and of mVCD to predict dementia over 5 years was investigated. RESULTS: The VASCOG criteria yielded rates of mVCD slightly lower than for DSM-5 and VICCCS. VaD rates were similar for all criteria, although slightly lower for DSM-IV. Agreement between the VASCOG, VICCCS and DSM-5 criteria was excellent for VaD and mVCD (κ = 0.83-1.0), but lower for VaD between VASCOG and the other criteria (κ = 0.47-0.63). VaD-based mortality predictions were similar for the VASCOG, VICCCS and DSM-5 criteria, and higher than those for other criteria. The prediction of incident dementia within 5 years from mVCD was slightly lower with VASCOG criteria than with DSM-5 and VICCCS criteria. CONCLUSIONS: The VASCOG criteria have greater sensitivity, modest concurrent validity and better predictive validity than older criteria for VaD, but are comparable to DSM-5 and VICCCS criteria. Their operationalization and inclusion of a mild VCD category make them useful for clinical and research applications.

Neural substrates for allocentric-to-egocentric conversion of remembered reach targets in humans.

Targets for goal-directed action can be encoded in allocentric coordinates (relative to another visual landmark), but it is not known how these are converted into egocentric commands for action. Here, we investigated this using a slow event-related fMRI paradigm, based on our previous behavioural finding that the allocentric-to-egocentric (Allo-Ego) conversion for reach is performed at the first possible opportunity. Participants were asked to remember (and eventually reach towards) the location of a briefly presented target relative to another visual landmark. After a first memory delay, participants were forewarned by a verbal instruction if the landmark would reappear at the same location (potentially allowing them to plan a reach following the auditory cue before the second delay), or at a different location where they had to wait for the final landmark to be presented before response, and then reach towards the remembered target location. As predicted, participants showed landmark-centred directional selectivity in occipital-temporal cortex during the first memory delay, and only developed egocentric directional selectivity in occipital-parietal cortex during the second delay for the 'Same cue' task, and during response for the 'Different cue' task. We then compared cortical activation between these two tasks at the times when the Allo-Ego conversion occurred, and found common activation in right precuneus, right presupplementary area and bilateral dorsal premotor cortex. These results confirm that the brain converts allocentric codes to egocentric plans at the first possible opportunity, and identify the four most likely candidate sites specific to the Allo-Ego transformation for reaches.

Temporal Evolution of Target Representation, Movement Direction Planning, and Reach Execution in Occipital-Parietal-Frontal Cortex: An fMRI Study.

The cortical mechanisms for reach have been studied extensively, but directionally selective mechanisms for visuospatial target memory, movement planning, and movement execution have not been clearly differentiated in the human. We used an event-related fMRI design with a visuospatial memory delay, followed by a pro-/anti-reach instruction, a planning delay, and finally a "go" instruction for movement. This sequence yielded temporally separable preparatory responses that expanded from modest parieto-frontal activation for visual target memory to broad occipital-parietal-frontal activation during planning and execution. Using the pro/anti instruction to differentiate visual and motor directional selectivity during planning, we found that one occipital area showed contralateral "visual" selectivity, whereas a broad constellation of left hemisphere occipital, parietal, and frontal areas showed contralateral "movement" selectivity. Temporal analysis of these areas through the entire memory-planning sequence revealed early visual selectivity in most areas, followed by movement selectivity in most areas, with all areas showing a stereotypical visuo-movement transition. Cross-correlation of these spatial parameters through time revealed separate spatiotemporally correlated modules for visual input, motor output, and visuo-movement transformations that spanned occipital, parietal, and frontal cortex. These results demonstrate a highly distributed occipital-parietal-frontal reach network involved in the transformation of retrospective sensory information into prospective movement plans.

Parietal area BA7 integrates motor programs for reaching, grasping, and bimanual coordination.

Skillful interaction with the world requires that the brain uses a multitude of sensorimotor programs and subroutines, such as for reaching, grasping, and the coordination of the two body halves. However, it is unclear how these programs operate together. Networks for reaching, grasping, and bimanual coordination might converge in common brain areas. For example, Brodmann area 7 (BA7) is known to activate in disparate tasks involving the three types of movements separately. Here, we asked whether BA7 plays a key role in integrating coordinated reach-to-grasp movements for both arms together. To test this, we applied transcranial magnetic stimulation (TMS) to disrupt BA7 activity in the left and right hemispheres, while human participants performed a bimanual size-perturbation grasping task using the index and middle fingers of both hands to grasp a rectangular object whose orientation (and thus grasp-relevant width dimension) might or might not change. We found that TMS of the right BA7 during object perturbation disrupted the bimanual grasp and transport/coordination components, and TMS over the left BA7 disrupted unimanual grasps. These results show that right BA7 is causally involved in the integration of reach-to-grasp movements of the two arms. NEW & NOTEWORTHY: Our manuscript describes a role of human Brodmann area 7 (BA7) in the integration of multiple visuomotor programs for reaching, grasping, and bimanual coordination. Our results are the first to suggest that right BA7 is critically involved in the coordination of reach-to-grasp movements of the two arms. The results complement previous reports of right-hemisphere lateralization for bimanual grasps.

Cervical dystonia: a neural integrator disorder.

Ocular motor neural integrators ensure that eyes are held steady in straight-ahead and eccentric positions of gaze. Abnormal function of the ocular motor neural integrator leads to centripetal drifts of the eyes with consequent gaze-evoked nystagmus. In 2002 a neural integrator, analogous to that in the ocular motor system, was proposed for the control of head movements. Recently, a counterpart of gaze-evoked eye nystagmus was identified for head movements; in which the head could not be held steady in eccentric positions on the trunk. These findings lead to a novel pathophysiological explanation in cervical dystonia, which proposed that the abnormalities of head movements stem from a malfunctioning head neural integrator, either intrinsically or as a result of impaired cerebellar, basal ganglia, or peripheral feedback. Here we briefly recapitulate the history of the neural integrator for eye movements, then further develop the idea of a neural integrator for head movements, and finally discuss its putative role in cervical dystonia. We hypothesize that changing the activity in an impaired head neural integrator, by modulating feedback, could treat dystonia.

Action relevance induces an attentional weighting of representations in visual working memory.

Information maintained in visual working memory (VWM) can be strategically weighted according to its task-relevance. This is typically studied by presenting cues during the maintenance interval, but under natural conditions, the importance of certain aspects of our visual environment is mostly determined by intended actions. We investigated whether representations in VWM are also weighted with respect to their potential action relevance. In a combined memory and movement task, participants memorized a number of items and performed a pointing movement during the maintenance interval. The test item in the memory task was subsequently presented either at the movement goal or at another location. We found that performance was better for test items presented at a location that corresponded to the movement goal than for test items presented at action-irrelevant locations. This effect was sensitive to the number of maintained items, suggesting that preferential maintenance of action relevant information becomes particularly important when the demand on VWM is high. We argue that weighting according to action relevance is mediated by the deployment of spatial attention to action goals, with representations spatially corresponding to the action goal benefitting from this attentional engagement. Performance was also better at locations next to the action goal than at locations farther away, indicating an attentional gradient spreading out from the action goal. We conclude that our actions continue to influence visual processing at the mnemonic level, ensuring preferential maintenance of information that is relevant for current behavioral goals.

Effect of allocentric landmarks on primate gaze behavior in a cue conflict task.

The relative contributions of egocentric versus allocentric cues on goal-directed behavior have been examined for reaches, but not saccades. Here, we used a cue conflict task to assess the effect of allocentric landmarks on gaze behavior. Two head-unrestrained macaques maintained central fixation while a target flashed in one of eight radial directions, set against a continuously present visual landmark (two horizontal/vertical lines spanning the visual field, intersecting at one of four oblique locations 11° from the target). After a 100-ms delay followed by a 100-ms mask, the landmark was displaced by 8° in one of eight radial directions. After a second delay (300-700 ms), the fixation point extinguished, signaling for a saccade toward the remembered target. When the landmark was stable, saccades showed a significant but small (mean 15%) pull toward the landmark intersection, and endpoint variability was significantly reduced. When the landmark was displaced, gaze endpoints shifted significantly, not toward the landmark, but partially (mean 25%) toward a virtual target displaced like the landmark. The landmark had a larger influence when it was closer to initial fixation, and when it shifted away from the target, especially in saccade direction. These findings suggest that internal representations of gaze targets are weighted between egocentric and allocentric cues, and this weighting is further modulated by specific spatial parameters.

Allocentric versus egocentric representation of remembered reach targets in human cortex.

The location of a remembered reach target can be encoded in egocentric and/or allocentric reference frames. Cortical mechanisms for egocentric reach are relatively well described, but the corresponding allocentric representations are essentially unknown. Here, we used an event-related fMRI design to distinguish human brain areas involved in these two types of representation. Our paradigm consisted of three tasks with identical stimulus display but different instructions: egocentric reach (remember absolute target location), allocentric reach (remember target location relative to a visual landmark), and a nonspatial control, color report (report color of target). During the delay phase (when only target location was specified), the egocentric and allocentric tasks elicited widely overlapping regions of cortical activity (relative to the control), but with higher activation in parietofrontal cortex for egocentric task and higher activation in early visual cortex for allocentric tasks. In addition, egocentric directional selectivity (target relative to gaze) was observed in the superior occipital gyrus and the inferior occipital gyrus, whereas allocentric directional selectivity (target relative to a visual landmark) was observed in the inferior temporal gyrus and inferior occipital gyrus. During the response phase (after movement direction had been specified either by reappearance of the visual landmark or a pro-/anti-reach instruction), the parietofrontal network resumed egocentric directional selectivity, showing higher activation for contralateral than ipsilateral reaches. These results show that allocentric and egocentric reach mechanisms use partially overlapping but different cortical substrates and that directional specification is different for target memory versus reach response.

Hand placement near the visual stimulus improves orientation selectivity in V2 neurons.

Often, the brain receives more sensory input than it can process simultaneously. Spatial attention helps overcome this limitation by preferentially processing input from a behaviorally-relevant location. Recent neuropsychological and psychophysical studies suggest that attention is deployed to near-hand space much like how the oculomotor system can deploy attention to an upcoming gaze position. Here we provide the first neuronal evidence that the presence of a nearby hand enhances orientation selectivity in early visual processing area V2. When the hand was placed outside the receptive field, responses to the preferred orientation were significantly enhanced without a corresponding significant increase at the orthogonal orientation. Consequently, there was also a significant sharpening of orientation tuning. In addition, the presence of the hand reduced neuronal response variability. These results indicate that attention is automatically deployed to the space around a hand, improving orientation selectivity. Importantly, this appears to be optimal for motor control of the hand, as opposed to oculomotor mechanisms which enhance responses without sharpening orientation selectivity. Effector-based mechanisms for visual enhancement thus support not only the spatiotemporal dissociation of gaze and reach, but also the optimization of vision for their separate requirements for guiding movements.