Statistical Learning of Frequent Distractor Locations in Visual Search Involves Regional Signal Suppression in Early Visual Cortex.

Observers can learn locations where salient distractors appear frequently to reduce potential interference-an effect attributed to better suppression of distractors at frequent locations. But how distractor suppression is implemented in the visual cortex and within the frontoparietal attention networks remains unclear. We used fMRI and a regional distractor-location learning paradigm with two types of distractors defined in either the same (orientation) or a different (color) dimension to the target to investigate this issue. fMRI results showed that BOLD signals in early visual cortex were significantly reduced for distractors (as well as targets) occurring at the frequent versus rare locations, mirroring behavioral patterns. This reduction was more robust with same-dimension distractors. Crucially, behavioral interference was correlated with distractor-evoked visual activity only for same- (but not different-) dimension distractors. Moreover, with different- (but not same-) dimension distractors, a color-processing area within the fusiform gyrus was activated more when a distractor was present in the rare region versus being absent and more with a distractor in the rare versus frequent locations. These results support statistical learning of frequent distractor locations involving regional suppression in early visual cortex and point to differential neural mechanisms of distractor handling with different- versus same-dimension distractors.

Simultaneous modeling of reaction times and brain dynamics in a spatial cueing task.

Understanding how brain activity translates into behavior is a grand challenge in neuroscientific research. Simultaneous computational modeling of both measures offers to address this question. The extension of the dynamic causal modeling (DCM) framework for blood oxygenation level-dependent (BOLD) responses to behavior (bDCM) constitutes such a modeling approach. However, only very few studies have employed and evaluated bDCM, and its application has been restricted to binary behavioral responses, limiting more general statements about its validity. This study used bDCM to model reaction times in a spatial attention task, which involved two separate runs with either horizontal or vertical stimulus configurations. We recorded fMRI data and reaction times (n= 26) and compared bDCM with classical DCM and a behavioral Rescorla-Wagner model using Bayesian model selection and goodness of fit statistics. Results indicate that bDCM performed equally well as classical DCM when modeling BOLD responses and as good as the Rescorla-Wagner model when modeling reaction times. Although our data revealed practical limitations of the current bDCM approach that warrant further investigation, we conclude that bDCM constitutes a promising method for investigating the link between brain activity and behavior.

Visual Size Processing in Early Visual Cortex Follows Lateral Occipital Cortex Involvement.

Neural activation in the early visual cortex (EVC) reflects the perceived rather than retinal size of stimuli, suggesting that feedback possibly from extrastriate regions modulates retinal size information in EVC. Meanwhile, the lateral occipital cortex (LOC) has been suggested to be critically involved in object size processing. To test for the potential contributions of feedback modulations on size representations in EVC, we investigated the dynamics of relevant processes using transcranial magnetic stimulation (TMS). Specifically, we briefly disrupted the neural activity of EVC and LOC at early, intermediate, and late time windows while participants performed size judgment tasks in either an illusory or neutral context. TMS over EVC and LOC allowed determining whether these two brain regions are relevant for generating phenomenological size impressions. Furthermore, the temporal order of TMS effects allowed inferences on the dynamics of information exchange between the two areas. Particularly, if feedback signals from LOC to EVC are crucial for generating altered size representations in EVC, then TMS effects over EVC should be observed simultaneously or later than the effects following LOC stimulation. The data from 20 humans (13 females) revealed that TMS over both EVC and LOC impaired illusory size perception. However, the strongest effects of TMS applied over EVC occurred later than those of LOC, supporting a functionally relevant feedback modulation from LOC to EVC for scaling size information. Our results suggest that context integration and the concomitant change of perceived size require LOC and result in modulating representations in EVC via recurrent processing.SIGNIFICANCE STATEMENT How we perceive an object's size is not entirely determined by its physical size or the size of its retinal representation but also the spatial context. Using transcranial magnetic stimulation, we investigated the role of the early visual cortex (EVC) and the higher-level visual area, lateral occipital cortex (LOC), known to be critically involved in object processing, in transforming an initial retinal representation into one that reflects perceived size. Transcranial magnetic stimulation altered size perception earlier over LOC compared with EVC, suggesting that context integration and the concomitant change in perceived size representations in EVC rely on feedback from LOC.

Feedback from lateral occipital cortex to V1/V2 triggers object completion: Evidence from functional magnetic resonance imaging and dynamic causal modeling.

Illusory figures demonstrate the visual system's ability to integrate disparate parts into coherent wholes. We probed this object integration process by either presenting an integrated diamond shape or a comparable ungrouped configuration that did not render a complete object. Two tasks were used that either required localization of a target dot (relative to the presented configuration) or discrimination of the dot's luminance. The results showed that only when the configuration was task relevant (in the localization task), performance benefited from the presentation of an integrated object. Concurrent functional magnetic resonance imaging was performed and analyzed using dynamic causal modeling to investigate the (causal) relationship between regions that are associated with illusory figure completion. We found object-specific feedback connections between the lateral occipital cortex (LOC) and early visual cortex (V1/V2). These modulatory connections persisted across task demands and hemispheres. Our results thus provide direct evidence that interactions between mid-level and early visual processing regions engage in illusory figure perception. These data suggest that LOC first integrates inputs from multiple neurons in lower-level cortices, generating a global shape representation while more fine-graded object details are then determined via feedback to early visual areas, independently of the current task demands.

Tracking the completion of parts into whole objects: Retinotopic activation in response to illusory figures in the lateral occipital complex.

Illusory figures demonstrate the visual system's ability to integrate separate parts into coherent, whole objects. The present study was performed to track the neuronal object construction process in human observers, by incrementally manipulating the grouping strength within a given configuration until the emergence of a whole-object representation. Two tasks were employed: First, in the spatial localization task, object completion could facilitate performance and was task-relevant, whereas it was irrelevant in the second, luminance discrimination task. Concurrent functional magnetic resonance imaging (fMRI) used spatial localizers to locate brain regions representing task-critical illusory-figure parts to investigate whether the step-wise object construction process would modulate neural activity in these localized brain regions. The results revealed that both V1 and the lateral occipital complex (LOC, with sub-regions LO1 and LO2) were involved in Kanizsa figure processing. However, completion-specific activations were found predominantly in LOC, where neural activity exhibited a modulation in accord with the configuration's grouping strength, whether or not the configuration was relevant to performing the task at hand. Moreover, right LOC activations were confined to LO2 and responded primarily to surface and shape completions, whereas left LOC exhibited activations in both LO1 and LO2 and was related to encoding shape structures with more detail. Together, these results demonstrate that various grouping properties within a visual scene are integrated automatically in LOC, with sub-regions located in different hemispheres specializing in the component sub-processes that render completed objects.

Attentional reorientation along the meridians of the visual field: Are there different neural mechanisms at play?

Hemispatial neglect, after unilateral lesions to parietal brain areas, is characterized by an inability to respond to unexpected stimuli in contralesional space. As the visual field's horizontal meridian is most severely affected, the brain networks controlling visuospatial processes might be tuned explicitly to this axis. We investigated such a potential directional tuning in the dorsal and ventral frontoparietal attention networks, with a particular focus on attentional reorientation. We used an orientation-discrimination task where a spatial precue indicated the target position with 80% validity. Healthy participants (n = 29) performed this task in two runs and were required to (re-)orient attention either only along the horizontal or the vertical meridian, while fMRI and behavioral measures were recorded. By using a general linear model for behavioral and fMRI data, dynamic causal modeling for effective connectivity, and other predictive approaches, we found strong statistical evidence for a reorientation effect for horizontal and vertical runs. However, neither neural nor behavioral measures differed between vertical and horizontal reorienting. Moreover, models from one run successfully predicted the cueing condition in the respective other run. Our results suggest that activations in the dorsal and ventral attention networks represent higher-order cognitive processes related to spatial attentional (re-)orientating that are independent of directional tuning and that unilateral attention deficits after brain damage are based on disrupted interactions between higher-level attention networks and sensory areas.

Combined expectancies: the role of expectations for the coding of salient bottom-up signals.

The visual system forms predictions about upcoming visual features based on previous visual experiences. Such predictions impact on current perception, so that expected stimuli can be detected faster and with higher accuracy. A key question is how these predictions are formed and on which levels of processing they arise. Particularly, predictions could be formed on early levels of processing, where visual features are represented separately, or might require higher levels of processing, with predictions formed based on full object representations that involve combinations of visual features. In four experiments, the present study investigated whether the visual system forms joint prediction errors or whether expectations about different visual features such as color and orientation are formed independently. The first experiment revealed that task-irrelevant and implicitly learned expectations were formed independently when the features were separately bound to different objects. In a second experiment, no evidence for a mutual influence of both types of task-irrelevant and implicitly formed feature expectations was observed, although both visual features were assigned to the same objects. A third experiment confirmed the findings of the previous experiments for explicitly rather than implicitly formed expectations. Finally, no evidence for a mutual influence of different feature expectations was observed when features were assigned to a single centrally presented object. Overall, the present results do not support the view that object feature binding generates joint feature-based expectancies of different object features. Rather, the results suggest that expectations for color and orientation are processed and resolved independently at the feature level.

Spatiotopic Adaptation in Visual Areas.

UNLABELLED: The ability to perceive the visual world around us as spatially stable despite frequent eye movements is one of the long-standing mysteries of neuroscience. The existence of neural mechanisms processing spatiotopic information is indispensable for a successful interaction with the external world. However, how the brain handles spatiotopic information remains a matter of debate. We here combined behavioral and fMRI adaptation to investigate the coding of spatiotopic information in the human brain. Subjects were adapted by a prolonged presentation of a tilted grating. Thereafter, they performed a saccade followed by the brief presentation of a probe. This procedure allowed dissociating adaptation aftereffects at retinal and spatiotopic positions. We found significant behavioral and functional adaptation in both retinal and spatiotopic positions, indicating information transfer into a spatiotopic coordinate system. The brain regions involved were located in ventral visual areas V3, V4, and VO. Our findings suggest that spatiotopic representations involved in maintaining visual stability are constructed by dynamically remapping visual feature information between retinotopic regions within early visual areas. SIGNIFICANCE STATEMENT: Why do we perceive the visual world as stable, although we constantly perform saccadic eye movements? We investigated how the visual system codes object locations in spatiotopic (i.e., external world) coordinates. We combined visual adaptation, in which the prolonged exposure to a specific visual feature alters perception, with fMRI adaptation, where the repeated presentation of a stimulus leads to a reduction in the BOLD amplitude. Functionally, adaptation was found in visual areas representing the retinal location of an adaptor but also at representations corresponding to its spatiotopic position. The results suggest that an active dynamic shift transports information in visual cortex to counteract the retinal displacement associated with saccade eye movements.

Neural correlates underlying the attentional spotlight in human parietal cortex independent of task difficulty.

Changes in the size of the attentional focus and task difficulty often co-vary. Nevertheless, the neural processes underlying the attentional spotlight process and task difficulty are likely to differ from each other. To differentiate between the two, we parametrically varied the size of the attentional focus in a novel behavioral paradigm while keeping visual processing difficulty either constant or not. A behavioral control experiment proved that the present behavioral paradigm could indeed effectively manipulate the size of the attentional focus per se, rather than affecting purely perceptual processes or surface processing. Imaging results showed that neural activity in a dorsal frontoparietal network, including right superior parietal cortex (SPL), was positively correlated with the size of the attentional spotlight, irrespective of whether task difficulty was constant or varied across different sizes of attentional focus. In contrast, neural activity in the ventral frontoparietal network, including the right inferior parietal cortex (IPL), was positively correlated with increasing task difficulty. Data suggest that sub-regions in parietal cortex are differentially involved in the attentional spotlight process and task difficulty: while SPL was involved in the attentional spotlight process independent of task difficulty, IPL was involved in the effect of task difficulty independent of the attentional spotlight process. Hum Brain Mapp 38:4996-5018, 2017. © 2017 Wiley Periodicals, Inc.

Spatiotopic updating of visual feature information.

Saccades shift the retina with high-speed motion. In order to compensate for the sudden displacement, the visuomotor system needs to combine saccade-related information and visual metrics. Many neurons in oculomotor but also in visual areas shift their receptive field shortly before the execution of a saccade (Duhamel, Colby, & Goldberg, 1992; Nakamura & Colby, 2002). These shifts supposedly enable the binding of information from before and after the saccade. It is a matter of current debate whether these shifts are merely location based (i.e., involve remapping of abstract spatial coordinates) or also comprise information about visual features. We have recently presented fMRI evidence for a feature-based remapping mechanism in visual areas V3, V4, and VO (Zimmermann, Weidner, Abdollahi, & Fink, 2016). In particular, we found fMRI adaptation in cortical regions representing a stimulus' retinotopic as well as its spatiotopic position. Here, we asked whether spatiotopic adaptation exists independently from retinotopic adaptation and which type of information is behaviorally more relevant after saccade execution. We first adapted at the saccade target location only and found a spatiotopic tilt aftereffect. Then, we simultaneously adapted both the fixation and the saccade target location but with opposite tilt orientations. As a result, adaptation from the fixation location was carried retinotopically to the saccade target position. The opposite tilt orientation at the retinotopic location altered the effects induced by spatiotopic adaptation. More precisely, it cancelled out spatiotopic adaptation at the saccade target location. We conclude that retinotopic and spatiotopic visual adaptation are independent effects.

The source of visual size adaptation.

Size adaptation describes the tendency of the visual system to adjust neural responsiveness of size representations after prolonged exposure to particular stimulations. A larger (or smaller) adaptor stimulus influences the perceived size of a similar test stimulus shown subsequently. Size adaptation may emerge on various processing levels. Functional representations of the adaptor to which the upcoming stimulus is adapted may be coded early in the visual system mainly reflecting retinal size. Alternatively, size adaptation may involve higher order processes that take into account additional information such as an object's estimated distance from the observer, hence reflecting perceived size. The present study investigated whether size adaptation is based on the retinal or the perceived size of an adaptor stimulus. A stimulus' physical and perceived sizes were orthogonally varied using perceived depth via binocular disparity, employing polarized 3D glasses. Four different adaptors were used, which varied in physical size, perceived size, or both. Two pairs of adaptors which were identical in physical size did not cause significantly different adaptation effects although they elicited different perceived sizes which were sufficiently large to produce differential aftereffects when induced by stimuli that physically differed in size. In contrast, there was a significant aftereffect when adaptors differed in physical size but were matched in perceived size. Size adaptation was thus unaffected by perceived size and binocular disparity. Our data suggest that size adaptation emerges from neural stages where information from both eyes is still coded in separate channels without binocular interactions, such as the lateral geniculate nucleus.

The Role of Top-Down Focused Spatial Attention in Preattentive Salience Coding and Salience-based Attentional Capture.

Selective visual attention requires an efficient coordination between top-down and bottom-up attention control mechanisms. This study investigated the behavioral and neural effects of top-down focused spatial attention on the coding of highly salient distractors and their tendency to capture attention. Combining spatial cueing with an irrelevant distractor paradigm revealed bottom-up based attentional capture only when attention was distributed across the whole search display, including the distractor location. Top-down focusing spatial attention on the target location abolished attentional capture of a salient distractor outside the current attentional focus. Functional data indicated that the missing capture effect was not based on diminished bottom-up salience signals at unattended distractor locations. Irrespectively of whether salient distractors occurred at attended or unattended locations, their presence enhanced BOLD signals at their respective spatial representation in early visual areas as well as in inferior frontal, superior parietal, and medial parietal cortex. Importantly, activity in these regions reflected the presence of a salient distractor rather than attentional capture per se. Moreover, successfully inhibiting attentional capture of a salient distractor at an unattended location further increased neural responses in medial parietal regions known to be involved in controlling spatial attentional shifts. Consequently, data provide evidence that top-down focused spatial attention prevents automatic attentional capture by supporting attentional control processes counteracting a spatial bias toward a salient distractor.

Individual attentional selection capacities are reflected in interhemispheric connectivity of the parietal cortex.

Modelling psychophysical data using the Theory of Visual Attention (TVA) allows for a quantification of attentional sub-processes, such as the resolution of competition amongst multiple stimuli by top-down control signals for target selection (TVA-parameter α). This fMRI study investigated the neural correlates of α by comparing activity differences and changes of effective connectivity between conditions where a target was accompanied by a distractor or by a second target. Twenty-five participants performed a partial report task inside the MRI scanner. The left angular gyrus (ANG), medial frontal, and posterior cingulate cortex showed higher activity when a target was accompanied by a distractor as opposed to a second target. The reverse contrast yielded activation of a bilateral fronto-parietal network, the anterior insula, anterior cingulate cortex, and left inferior occipital gyrus. A psychophysiological interaction analysis revealed that the connectivity between left ANG and the left and right supramarginal gyrus (SMG), left anterior insula, and right putamen was enhanced in the target-distractor condition in participants with worse attentional top-down control. Dynamic causal modelling suggested that the connection from left ANG to right SMG during distractor presence was modulated by α. Our data show that interindividual differences in attentional processing are reflected in changes of effective connectivity without significant differences in activation strength of network nodes.

Rescaling retinal size into perceived size: evidence for an occipital and parietal bottleneck.

The spatial and temporal context of an object influence its perceived size. Two visual illusions illustrate this nicely: the size adaptation effect and the Ebbinghaus illusion. Whereas size adaptation affects size rescaling of a target circle via a previously presented, differently sized adaptor circle, the Ebbinghaus illusion alters perceived size by virtue of surrounding circles. In the classical Ebbinghaus setting, the surrounding circles are shown simultaneously with the target. However, size underestimation persists when the surrounding circles precede the target. Such a temporal separation of inducer and target circles in both illusions permits the comparison of BOLD signals elicited by two displays that, although objectively identical, elicit different percepts. The current study combined both illusions in a factorial design to identify a presumed common central mechanism involved in rescaling retinal into perceived size. At the behavioral level, combining both illusions did not affect perceived size further. At the neural level, however, this combination induced functional activation beyond that induced by either illusion separately: An underadditive activation pattern was found within left lingual gyrus, right supramarginal gyrus, and right superior parietal cortex. These findings provide direct behavioral and functional evidence for the presence of a neural bottleneck in rescaling retinal into perceived size, a process vital for visual perception.

Selecting category specific visual information: Top-down and bottom-up control of object based attention.

The ability to select, within the complexity of sensory input, the information most relevant for our purposes is influenced by both internal settings (i.e., top-down control) and salient features of external stimuli (i.e., bottom-up control). We here investigated using fMRI the neural underpinning of the interaction of top-down and bottom-up processes, as well as their effects on extrastriate areas processing visual stimuli in a category-selective fashion. We presented photos of bodies or buildings embedded into frequency-matched visual noise to the subjects. Stimulus saliency changed gradually due to an altered degree to which photos stood-out in relation to the surrounding noise (hence generating stronger bottom-up control signals). Top-down settings were manipulated via instruction: participants were asked to attend one stimulus category (i.e., "is there a body?" or "is there a building?"). Highly salient stimuli that were inconsistent with participants' attentional top-down template activated the inferior frontal junction and dorsal parietal regions bilaterally. Stimuli consistent with participants' current attentional set additionally activated insular cortex and the parietal operculum. Furthermore, the extrastriate body area (EBA) exhibited increased neural activity when attention was directed to bodies. However, the latter effect was found only when stimuli were presented at intermediate saliency levels, thus suggesting a top-down modulation of this region only in the presence of weak bottom-up signals. Taken together, our results highlight the role of the inferior frontal junction and posterior parietal regions in integrating bottom-up and top-down attentional control signals.

Attention modulates visual size adaptation.

The current study determined in healthy subjects (n = 16) whether size adaptation occurs at early, i.e., preattentive, levels of processing or whether higher cognitive processes such as attention can modulate the illusion. To investigate this issue, bottom-up stimulation was kept constant across conditions by using a single adaptation display containing both small and large adapter stimuli. Subjects' attention was directed to either the large or small adapter stimulus by means of a luminance detection task. When attention was directed toward the small as compared to the large adapter, the perceived size of the subsequent target was significantly increased. Data suggest that different size adaptation effects can be induced by one and the same stimulus depending on the current allocation of attention. This indicates that size adaptation is subject to attentional modulation. These findings are in line with previous research showing that transient as well as sustained attention modulates visual features, such as contrast sensitivity and spatial frequency, and influences adaptation in other contexts, such as motion adaptation (Alais & Blake, 1999; Lankheet & Verstraten, 1995). Based on a recently suggested model (Pooresmaeili, Arrighi, Biagi, & Morrone, 2013), according to which perceptual adaptation is based on local excitation and inhibition in V1, we conclude that guiding attention can boost these local processes in one or the other direction by increasing the weight of the attended adapter. In sum, perceptual adaptation, although reflected in changes of neural activity at early levels (as shown in the aforementioned study), is nevertheless subject to higher-order modulation.

The moon illusion and size-distance scaling--evidence for shared neural patterns.

A moon near to the horizon is perceived larger than a moon at the zenith, although--obviously--the moon does not change its size. In this study, the neural mechanisms underlying the "moon illusion" were investigated using a virtual 3-D environment and fMRI. Illusory perception of an increased moon size was associated with increased neural activity in ventral visual pathway areas including the lingual and fusiform gyri. The functional role of these areas was further explored in a second experiment. Left V3v was found to be involved in integrating retinal size and distance information, thus indicating that the brain regions that dynamically integrate retinal size and distance play a key role in generating the moon illusion.

Modulation of top-down control of visual attention by cathodal tDCS over right IPS.

The right intraparietal sulcus (rIPS) is a key region for the endogenous control of selective visual attention in the human brain. Previous studies suggest that the rIPS is especially involved in top-down control and spatial distribution of attention across both visual hemifields. We further explored these attentional functions using transcranial direct current stimulation (tDCS) of the rIPS to modulate behavioral performance in a partial report task. Performance was analyzed according to the theory of visual attention (TVA) (Bundesen, 1990), which provides a computational framework to investigate different parameters of visuo-attentional processing such as top-down control, attentional weighting, capacity of visual short term memory, and processing speed. We investigated the effects of different tDCS current strengths (1 mA and 2 mA) in two experiments: 1 mA tDCS (anodal, cathodal, sham) did not affect any of the TVA parameters, but cathodal 2 mA stimulation significantly enhanced top-down control as evidenced by a reduction of the α parameter of TVA, regardless of hemifield. This differential impact on the top-down control component of attentional processing suggests that the horizontal rIPS is mainly involved in attentional selection as none of the spatial or resource variables of TVA were altered. Furthermore, the data add evidence to previous work highlighting (1) the importance of using appropriate current strength in stimulation protocols, and (2) that the often reported inhibitory effect of cathodal stimulation in e.g., motor tasks might not extend to cognitive paradigms.

Deconstructing the architecture of dorsal and ventral attention systems with dynamic causal modeling.

Attentional orientation to a spatial cue and reorientation-after invalid cueing-are mediated by two distinct networks in the human brain. A bilateral dorsal frontoparietal network, comprising the intraparietal sulcus (IPS) and the frontal eye fields (FEF), controls the voluntary deployment of attention and may modulate visual cortex in preparation for upcoming stimulation. In contrast, reorienting attention to invalidly cued targets engages a right-lateralized ventral frontoparietal network comprising the temporoparietal junction (TPJ) and ventral frontal cortex. The present fMRI study investigated the functional architecture of these two attentional systems by characterizing effective connectivity during lateralized orienting and reorienting of attention, respectively. Subjects performed a modified version of Posner's location-cueing paradigm. Dynamic causal modeling (DCM) of regional responses in the dorsal and ventral network, identified in a conventional (SPM) whole-brain analysis, was used to compare different functional architectures. Bayesian model selection showed that top-down connections from left and right IPS to left and right visual cortex, respectively, were modulated by the direction of attention. Moreover, model evidence was highest for a model with directed influences from bilateral IPS to FEF, and reciprocal coupling between right and left FEF. Invalid cueing enhanced forward connections from visual areas to right TPJ, and directed influences from right TPJ to right IPS and IFG (inferior frontal gyrus). These findings shed further light on the functional organization of the dorsal and ventral attentional network and support a context-sensitive lateralization in the top-down (backward) mediation of attentional orienting and the bottom-up (forward) effects of invalid cueing.

Neural mechanisms of attentional reorienting in three-dimensional space.

How the human brain reconstructs the three-dimensional (3D) world from two-dimensional (2D) retinal images has received a great deal of interest as has how we shift attention in 2D space. In contrast, it remains poorly understood how visuospatial attention is shifted in depth. In this fMRI study, by constructing a virtual 3D environment in the MR scanner and by presenting targets either close to or far from the participants in an adapted version of the Posner spatial-cueing paradigm, we investigated the behavioral and neural mechanisms underlying visuospatial orienting/reorienting in depth. At the behavioral level, although covering the same spatial distance, attentional reorienting to objects unexpectedly appearing closer to the observer and in the unattended hemispace was faster than reorienting to unexpected objects farther away. At the neural level, we found that in addition to the classical attentional reorienting system in the right temporoparietal junction, two additional brain networks were differentially involved in aspects of attentional reorienting in depth. First, bilateral premotor cortex reoriented visuospatial attention specifically along the third dimension of visual space (i.e., from close to far or vice versa), compared with attentional reorienting within the same depth plane. Second, a network of areas reminiscent of the human "default-mode network," including posterior cingulate cortex, orbital prefrontal cortex, and left angular gyrus, was involved in the neural interaction between depth and attentional orienting, by boosting attentional reorienting to unexpected objects appearing both closer to the observer and in the unattended hemispace.