Inferior Intraparietal Sulcus Research Articles

Causal evidence for distinct mechanisms of prioritization and suppression in the control of working memory

Abstract Working memory requires maintenance of relevant information and suppression of irrelevant information. Our previous study administered rhythmic transcranial magnetic stimulation (TMS) to anterior middle frontal gyrus and inferior intraparietal sulcus and found causal evidence for posterior alpha oscillations and anterior theta oscillation in the control over working memory. However, this previous study did not measure neural oscillations using electrophysiology. To investigate the specific neural mechanism for prioritization and suppression, we conducted a pre-registered four-session experiment with 47 participants that administered rhythmic TMS concurrent with electroencephalography (EEG) as participants performed a retro-cue working memory task. Rhythmic TMS was personalized to the peak frequency of theta and alpha oscillatory activity to increase its efficacy. Our baseline analysis revealed the canonical alpha lateralization pattern in posterior cortex to a retro-cue that indicated which visual hemifield would be tested at probe. Alpha lateralization was positively correlated with the benefit to working memory capacity from the retro-cue. In addition, we found that theta-frequency functional connectivity between the left frontal and parietal cortex increased to the retro-cue. During the stimulation sessions, we found evidence of target engagement such that theta-frequency TMS to frontal cortex and alpha-frequency TMS to parietal cortex successfully increased the amplitude of the targeted oscillation. We replicated our previous behavioral interaction between site and frequency such that rhythmic TMS improved working memory capacity when matched to endogenous activity. Finally, alpha-frequency TMS increased posterior alpha lateralization and theta-frequency TMS increased frontal-parietal theta-connectivity. Altogether, these results provide causal evidence for a model in which alpha lateralization supports suppression and frontal-parietal theta-connectivity supports prioritization. Keywords: Working memory, Rhythmic TMS, Concurrent TMS-EEG, Functional Connectivity

The role of the l-IPS in the comprehension of reversible and irreversible sentences: an rTMS study

Thematic roles can be seen as semantic labels assigned to who/what is taking part in the event denoted by a verb. Encoding thematic relations is crucial for sentence interpretation since it relies on both syntactic and semantic aspects. In previous studies, repetitive transcranial magnetic stimulation (rTMS) over the left inferior intraparietal sulcus (l-IPS) selectively influenced performance accuracy on reversible passive (but not active) sentences. The effect was attributed to the fact that in these sentences the assignment of the agent and theme roles requires re-analysis of the first-pass sentence parsing.To evaluate the role of reversibility and non-canonical word order (passive voice) on the effect, rTMS was applied over l-IPS during a sentence comprehension task that included reversible and irreversible, active and passive sentences. Participants were asked to identify who/what was performing the action or who/what the action was being performed on.Stimulation of the l-IPS increased response time on reversible passive sentences but not on reversible active sentences. Importantly, no effect was found on irreversible sentences, irrespective of sentence diathesis.Results suggest that neither reversibility nor sentence diathesis alone are responsible for the effect and that the effect is likely to be triggered/constrained by a combination of semantic reversibility and non-canonical word order. Combined with the results of previous studies, and irrespective of the specific role of each feature, these findings support the view that the l-IPS is critically involved in the assignment of thematic roles in reversible sentences.

Functional dissociation of anterior cingulate cortex and intraparietal sulcus in visual working memory

Previous electrophysiological studies of lateralized visual working memory (VWM) identified an ERP component, defined as contralateral delay activity (CDA), directly modulated by the number of items held in memory. One of the main candidate as the cortical source of this ERP component is the inferior intraparietal sulcus (IPS). Moreover, previous neuroimaging studies put forth evidence for the presence of a distributed VWM network involving also prefrontal areas and in particular the anterior cingulate cortex (ACC). Nonetheless, the understanding of the functional role of ACC is still debated. We recorded the high-density EEG in 20 healthy participants undergoing a VWM and a control task. Explorative cluster-based permutation statistics confirmed the posterior memory load dependent CDA modulation, but also identified an additional anterior cluster of electrodes whose amplitude was modulated by memory load. The source reconstruction revealed a memory load dependent activation in the IPS but also in the ACC, suggesting that these two areas might be nodes of a fronto-parietal circuit underlying VWM maintenance. Crucially, parietal and prefrontal areas showed a temporal dissociation, since IPS was more engaged in the early phase of visual information storage while the ACC was more active during the late phase. This pattern suggests a functional dissociation between the parietal cortex, which is involved in encoding and storage of information, and prefrontal areas, subserving cognitive control processes, including the boosting and protection of information from decay. Remarkably, the connection strength between IPS and ACC predicted the individual number of items held in memory. These findings are discussed within the theoretical account of a neural distributed model of VWM.

Individuation of parts of a single object and multiple distinct objects relies on a common neural mechanism in inferior intraparietal sulcus

Object identification and enumeration rely on the ability to distinguish, or individuate, objects from the background. Does multiple object individuation operate only over bounded, separable objects or does it operate equally over connected features within a single object? While previous fMRI experiments suggest that connectedness affects the processing and enumeration of objects, recent behavioral and EEG studies demonstrated that parallel individuation occurs over both object parts and distinct objects. However, it is unclear whether individuation of object parts and distinct objects relies on a common or independent neural mechanisms. Using fMRI-based multivariate pattern analyses, we here demonstrate that activity patterns in inferior and superior intraparietal sulci (IPS) encode numerosity independently of whether the individuated items are connected parts of a single object or distinct objects. Lateral occipital cortex is more sensitive to perceptual aspects of the two stimulus types and the targets of the stimuli, suggesting a dissociation between ventral and dorsal areas in representing perceptual object properties and more general information about numerosity, respectively. Our results suggest that objecthood is not a necessary prerequisite for parallel individuation in IPS. Rather, our results point toward a common individuation mechanism that selects targets over a flexible object hierarchy, independently of whether the targets are distinct separable objects or parts of a single object.

Neural activity during working memory encoding, maintenance, and retrieval: A network-based model and meta-analysis.

It remains unclear whether and to what extent working memory (WM) temporal subprocesses (i.e., encoding, maintenance, and retrieval) involve shared or distinct intrinsic networks. To address this issue, I constructed a model of intrinsic network contributions to different WM phases and then evaluated the validity of the model by performing a quantitative meta-analysis of relevant functional neuroimaging data. The model suggests that the transition from the encoding to maintenance and to retrieval stages involves progressively decreasing involvement of the dorsal attention network (DAN), but progressively increasing involvement of the frontoparietal control network (FPCN). Separate meta-analysis of each phase effect and direct comparisons between them yielded results that were largely consistent with the model. This evidence included between-phase double dissociations that were consistent with the model, such as encoding > maintenance contrast showing some DAN, but no FPCN, regions, and maintenance > encoding contrast showing the reverse, that is, some FPCN, but no DAN, regions. Two closely juxtaposed regions that are members of the DAN and FPCN, such as inferior frontal junction versus caudal prefrontal cortex and superior versus inferior intraparietal sulcus, showed a high degree of functional differentiation. Although all regions identified in the present study were already identified in previous WM studies, this study uniquely enhances our understating of their roles by clarifying their network membership and specific associations with different WM phases.

Interference during the retrieval of arithmetic and lexico-semantic knowledge modulates similar brain regions: Evidence from functional magnetic resonance imaging (fMRI)

Single-digit multiplications are mainly solved by memory retrieval. However, these problems are also prone to errors due to systematic interference (i.e., co-activation of interconnected but incorrect solutions). Semantic control processes are crucial to overcome this type of interference and to retrieve the correct information. Previous research suggests the importance of several brain regions such as the left inferior frontal cortex and the intraparietal sulcus (IPS) for semantic control. But, this evidence is mainly based on tasks measuring interference during the processing of lexico-semantic information (e.g., pictures or words). Here, we investigated whether semantic control during arithmetic problem solving (i.e., multiplication fact retrieval) draws upon similar or different brain mechanisms as in other semantic domains (i.e., lexico-semantic).The brain activity of 46 students was measured with fMRI while participants performed an operand-related-lure (OR) and a picture-word (PW) task. In the OR task participants had to verify the correctness of a given solution to a single-digit multiplication. Similarly, in the PW task, participants had to judge whether a presented word matches the concept displayed in a picture or not.Analyses showed that resolving interference in these two tasks modulates the activation of a widespread fronto-parietal network (e.g., left/right IFG, left insula lobe, left IPS). Importantly, conjunction analysis revealed a neural overlap in the left inferior frontal gyrus (IFG) pars triangularis and left IPS. Additional Bayesian analyses showed that regions that are thought to store lexico-semantic information (e.g., left middle temporal gyrus) did not show evidence for an arithmetic interference effect. Overall, our findings not only indicate that semantic control plays an important role in arithmetic problem solving but also that it is supported by common brain regions across semantic domains. Additionally, by conducting Bayesian analysis we confirmed the hypothesis that the semantic control network contributes differently to semantic tasks of various domains.

On the Role of the Inferior Intraparietal Sulcus in Visual Working Memory for Lateralized Single-feature Objects

A consolidated practice in cognitive neuroscience is to explore the properties of human visual working memory through the analysis of electromagnetic signals using cued change detection tasks. Under these conditions, EEG/MEG activity increments in the posterior parietal cortex scaling with the number of memoranda are often reported in the hemisphere contralateral to the objects' position in the memory array. This highly replicable finding clashes with several reported failures to observe compatible hemodynamic activity modulations using fMRI or fNIRS in comparable tasks. Here, we reconcile this apparent discrepancy by acquiring fMRI data on healthy participants and employing a cluster analysis to group voxels in the posterior parietal cortex based on their functional response. The analysis identified two distinct subpopulations of voxels in the intraparietal sulcus (IPS) showing a consistent functional response among participants. One subpopulation, located in the superior IPS, showed a bilateral response to the number of objects coded in visual working memory. A different subpopulation, located in the inferior IPS, showed an increased unilateral response when the objects were displayed contralaterally. The results suggest that a cluster of neurons in the inferior IPS is a candidate source of electromagnetic contralateral responses to working memory load in cued change detection tasks.

The impact of top-down spatial attention on laterality and hemispheric asymmetry in the human parietal cortex.

The human parietal cortex exhibits a preference to contralaterally presented visual stimuli (i.e., laterality) as well as an asymmetry between the two hemispheres with the left parietal cortex showing greater laterality than the right. Using visual short-term memory and perceptual tasks and varying target location predictability, this study examined whether hemispheric laterality and asymmetry are fixed characteristics of the human parietal cortex or whether they are dynamic and modulated by the deployment of top-down attention to the target present hemifield. Two parietal regions were examined here that have previously been shown to be involved in visual object individuation and identification and are located in the inferior and superior intraparietal sulcus (IPS), respectively. Across three experiments, significant laterality was found in both parietal regions regardless of attentional modulation with laterality being greater in the inferior than superior IPS, consistent with their roles in object individuation and identification, respectively. Although the deployment of top-down attention had no effect on the superior IPS, it significantly increased laterality in the inferior IPS. The deployment of top-down spatial attention can thus amplify the strength of laterality in the inferior IPS. Hemispheric asymmetry, on the other hand, was absent in both brain regions and only emerged in the inferior but not the superior IPS with the deployment of top-down attention. Interestingly, the strength of hemispheric asymmetry significantly correlated with the strength of laterality in the inferior IPS. Hemispheric asymmetry thus seems to only emerge when there is a sufficient amount of laterality present in a brain region.

Understanding location- and feature-based processing along the human intraparietal sulcus.

Based on different cognitive tasks and mapping methods, the human intraparietal sulcus (IPS) has been subdivided according to multiple different organizational schemes. The presence of topographically organized regions throughout IPS indicates a strong location-based processing in this brain region. However, visual short-term memory (VSTM) studies have shown that while a region in the inferior IPS region (inferior IPS) is involved in object individuation and selection based on location, a region in the superior IPS (superior IPS) primarily encodes and stores object featural information. Here, we determined the localization of these two VSTM IPS regions with respect to the topographic IPS regions in individual participants and the role of different IPS regions in location- and feature-based processing. Anatomically, inferior IPS showed an 85.2% overlap with topographic IPS regions, with the greatest overlap seen in V3A and V3B, and superior IPS showed a 73.6% overall overlap, with the greatest overlap seen in IPS0-2. Functionally, there appeared to be a partial overlap between IPS regions involved in location- and feature-based processing, with more inferior and medial regions showing a stronger location-based processing and more superior and lateral regions showing a stronger feature-based processing. Together, these results suggest that understanding the multiplex nature of IPS in visual cognition may not be reduced to examining the functions of the different IPS topographic regions, but rather, it can only be accomplished by understanding how regions identified by different tasks and methods may colocalize with each other.

The compensatory dynamic of inter-hemispheric interactions in visuospatial attention revealed using rTMS and fMRI.

A balance of mutual tonic inhibition between bi-hemispheric posterior parietal cortices is believed to play an important role in bilateral visual attention. However, experimental support for this notion has been mainly drawn from clinical models of unilateral damage. We have previously shown that low-frequency repetitive TMS (rTMS) over the intraparietal sulcus (IPS) generates a contralateral attentional deficit in bilateral visual tracking. Here, we used functional magnetic resonance imaging (fMRI) to study whether rTMS temporarily disrupts the inter-hemispheric balance between bilateral IPS in visual attention. Following application of 1 Hz rTMS over the left IPS, subjects performed a bilateral visual tracking task while their brain activity was recorded using fMRI. Behaviorally, tracking accuracy was reduced immediately following rTMS. Areas ventro-lateral to left IPS, including inferior parietal lobule (IPL), lateral IPS (LIPS), and middle occipital gyrus (MoG), showed decreased activity following rTMS, while dorsomedial areas, such as Superior Parietal Lobule (SPL), Superior occipital gyrus (SoG), and lingual gyrus, as well as middle temporal areas (MT+), showed higher activity. The brain activity of the homologues of these regions in the un-stimulated, right hemisphere was reversed. Interestingly, the evolution of network-wide activation related to attentional behavior following rTMS showed that activation of most occipital synergists adaptively compensated for contralateral and ipsilateral decrement after rTMS, while activation of parietal synergists, and SoG remained competing. This pattern of ipsilateral and contralateral activations empirically supports the hypothesized loss of inter-hemispheric balance that underlies clinical manifestation of visual attentional extinction.

The Role of Transverse Occipital Sulcus in Scene Perception and Its Relationship to Object Individuation in Inferior Intraparietal Sulcus

The parietal cortex has been functionally divided into various subregions; however, very little is known about how these areas relate to each other. Two such regions are the transverse occipital sulcus (TOS) scene area and inferior intraparietal sulcus (IPS). TOS exhibits similar activation patterns to the scene selective parahippocampal place area, suggesting its role in scene perception. Inferior IPS, in contrast, has been shown to participate in object individuation and selection via location. Interestingly, both regions have been localized to the same general area of the brain. If these two were actually the same brain region, it would have important implications regarding these regions' role in cognition. To explore this, we first localized TOS and inferior IPS in individual participants and examined the degree of overlap between these regions in each participant. We found that TOS showed only a minor degree of overlap with inferior IPS (∼10%). We then directly explored the role of TOS and inferior IPS in object individuation and scene perception by examining their responses to furnished rooms, empty rooms, isolated furniture, and multiple isolated objects. If TOS and inferior IPS were the same region, we would expect to see similar response patterns in both. Instead, the response of TOS was predominantly scene selective, whereas activity in inferior IPS was primarily driven by the number of objects present in the display, regardless of scene context. These results show that TOS and inferior IPS are nearby but distinct regions, with different functional roles in visual cognition.

Neural Representation of Targets and Distractors during Object Individuation and Identification

In many everyday activities, we need to attend and encode multiple target objects among distractor objects. For example, when driving a car on a busy street, we need to simultaneously attend objects such as traffic signs, pedestrians, and other cars, while ignoring colorful and flashing objects in display windows. To explain how multiple visual objects are selected and encoded in visual STM and in perception in general, the neural object file theory argues that, whereas object selection and individuation is supported by inferior intraparietal sulcus (IPS), the encoding of detailed object features that enables object identification is mediated by superior IPS and higher visual areas such as the lateral occipital complex (LOC). Nevertheless, because task-irrelevant distractor objects were never present in previous studies, it is unclear how distractor objects would impact neural responses related to target object individuation and identification. To address this question, in two fMRI experiments, we asked participants to encode target object shapes among distractor object shapes, with targets and distractors shown in different spatial locations and in different colors. We found that distractor-related neural processing only occurred at low, but not at high, target encoding load and impacted both target individuation in inferior IPS and target identification in superior IPS and LOC. However, such distractor-related neural processing was short-lived, as it was only present during the visual STM encoding but not the delay period. Moreover, with spatial cuing of target locations in advance, distractor processing was attenuated during target encoding in superior IPS. These results are consistent with the load theory of visual information processing. They also show that, whereas inferior IPS and LOC were automatically engaged in distractor processing under low task load, with the help of precuing, superior IPS was able to only encode the task-relevant visual information.

Visual Working Memory in Human Cortex

Visual working memory (VWM) is the ability to maintain visual information in a readily available and easily updated state. Converging evidence has revealed that VWM capacity is limited by the number of maintained objects, which is about 3 - 4 for the average human. Recent work suggests that VWM capacity is also limited by the resolution required to maintain objects, which is tied to the objects' inherent complexity. Electroencephalogram (EEG) studies using the Contralateral Delay Activity (CDA) paradigm have revealed that cortical representations of VWM are at a minimum loosely organized like the primary visual system, such that the left side of space is represented in the right hemisphere, and vice versa. Recent functional magnetic resonance imaging (fMRI) work shows that the number of objects is maintained by representations in the inferior intraparietal sulcus (IPS) along dorsal parietal cortex, whereas the resolution of these maintained objects is subserved by the superior IPS and the lateral occipital complex (LOC). These areas overlap with recently-discovered, retinotopically-organized visual field maps (VFMs) spanning the IPS (IPS-0/1/2/3/4/5), and potentially maps in lateral occipital cortex, such as LO-1/2, and/or TO-1/2 (hMT+). Other fMRI studies have implicated early VFMs in posterior occipital cortex, suggesting that visual areas V1-hV4 are recruited to represent information in VWM. Insight into whether and how these VFMs subserve VWM may illuminate the nature of VWM. In addition, understanding the nature of these maps may allow a greater investigation into individual differences among subjects and even between hemispheres within subjects.

Brain networks of novelty-driven involuntary and cued voluntary auditory attention shifting.

In everyday life, we need a capacity to flexibly shift attention between alternative sound sources. However, relatively little work has been done to elucidate the mechanisms of attention shifting in the auditory domain. Here, we used a mixed event-related/sparse-sampling fMRI approach to investigate this essential cognitive function. In each 10-sec trial, subjects were instructed to wait for an auditory “cue” signaling the location where a subsequent “target” sound was likely to be presented. The target was occasionally replaced by an unexpected “novel” sound in the uncued ear, to trigger involuntary attention shifting. To maximize the attention effects, cues, targets, and novels were embedded within dichotic 800-Hz vs. 1500-Hz pure-tone “standard” trains. The sound of clustered fMRI acquisition (starting at t = 7.82 sec) served as a controlled trial-end signal. Our approach revealed notable activation differences between the conditions. Cued voluntary attention shifting activated the superior intra­­parietal sulcus (IPS), whereas novelty-triggered involuntary orienting activated the inferior IPS and certain subareas of the precuneus. Clearly more widespread activations were observed during voluntary than involuntary orienting in the premotor cortex, including the frontal eye fields. Moreover, we found ­evidence for a frontoinsular-cingular attentional control network, consisting of the anterior insula, inferior frontal cortex, and medial frontal cortices, which were activated during both target discrimination and voluntary attention shifting. Finally, novels and targets activated much wider areas of superior temporal auditory cortices than shifting cues.

A computational model of fMRI activity in the intraparietal sulcus that supports visual working memory

A computational model was developed to explain a pattern of results of fMRI activation in the intraparietal sulcus (IPS) supporting visual working memory for multiobject scenes. The model is based on the hypothesis that dendrites of excitatory neurons are major computational elements in the cortical circuit. Dendrites enable formation of a competitive queue that exhibits a gradient of activity values for nodes encoding different objects, and this pattern is stored in working memory. In the model, brain imaging data are interpreted as a consequence of blood flow arising from dendritic processing. Computer simulations showed that the model successfully simulates data showing the involvement of inferior IPS in object individuation and spatial grouping through representation of objects' locations in space, along with the involvement of superior IPS in object identification through representation of a set of objects' features. The model exhibits a capacity limit due to the limited dynamic range for nodes and the operation of lateral inhibition among them. The capacity limit is fixed in the inferior IPS regardless of the objects' complexity, due to the normalization of lateral inhibition, and variable in the superior IPS, due to the different encoding demands for simple and complex shapes. Systematic variation in the strength of self-excitation enables an understanding of the individual differences in working memory capacity. The model offers several testable predictions regarding the neural basis of visual working memory.

The neural mechanisms of visual selection: the view from neuropsychology

In this review, we discuss how neuropsychological impairments in visual selection can inform us about how selection normally operates. Using neuroanatomical and behavioral evidence on the disorders of neglect, extinction, and simultanagnosia, we propose functional and anatomical links between different aspects of visual selection and distinct sites in the posterior parietal cortex (PPC). This includes linking: (i) bottom-up attentional capture and the right temporo-parietal junction (TPJ); (ii) top-down segmentation of displays and the medial PPC; (iii) grouping, individuation and identification, and the inferior intra-parietal sulcus (IPS) bilaterally; and (iv) the suppression of saliency and the left IPS. In addition, when neuropsychological studies are combined with fMRI, there is evidence that these regions of the PPC interact with striate and extra-striate cortical areas, which respond to specific properties of stimuli. Selection should be viewed as an emergent property of a network of areas involving both ventral and dorsal cortex.

Discrete Object Representation, Attention Switching, and Task Difficulty in the Parietal Lobe

An important component of perception, attention, and memory is the structuring of information into subsets ("objects"), which allows some parts to be considered together but kept separate from others. Portions of the posterior parietal lobe respond proportionally to the number of objects in the scope of attention and short-term memory, up to a capacity limit of around four, suggesting they have a role in this important process. This study investigates the relationship of discrete object representation to other parietal functions. Two experiments and two supplementary analyses were conducted to evaluate responsivity in parietal regions to the number of objects, the number of spatial locations, attention switching, and general task difficulty. Using transparent motion, it was found that a posterior and inferior parietal response to multiple objects persists even in the absence of a change in visual extent or the number of spatial locations. In a monitoring task, it was found that attention switching (or task difficulty) and object representation have distinct neural signatures, with the former showing greater recruitment of an anterior and lateral intraparietal sulcus (IPS) region, but the latter in a posterior and lateral region. A dissociation was also seen between selectivity for object load across tasks in the inferior IPS and feature or object-related memory load in the superior IPS.

Oscillatory activity in parietal and dorsolateral prefrontal cortex during retention in visual short‐term memory: Additive effects of spatial attention and memory load

We used whole-head magnetoencephalography to study the representation of objects in visual short-term memory (VSTM) in the human brain. Subjects remembered the location and color of either two or four colored disks that were encoded from the left or right visual field (equal number of distractors in the other visual hemifield). The data were analyzed using time-frequency methods, which enabled us to discover a strong oscillatory activity in the 8-15 Hz band during the retention interval. The study of the alpha power variation revealed two types of responses, in different brain regions. The first was a decrease in alpha power in parietal cortex, contralateral to the stimuli, with no load effect. The second was an increase of alpha power in parietal and lateral prefrontal cortex, as memory load increased, but without interaction with the hemifield of the encoded stimuli. The absence of interaction between side of encoded stimuli and memory load suggests that these effects reflect distinct underlying mechanisms. A novel method to localize the neural generators of load-related oscillatory activity was devised, using cortically-constrained distributed source-localization methods. Some activations were found in the inferior intraparietal sulcus (IPS) and intraoccipital sulcus (IOS). Importantly, strong oscillatory activity was also found in dorsolateral prefrontal cortex (DLPFC). Alpha oscillatory activity in DLPFC was synchronized with the activity in parietal regions, suggesting that VSTM functions in the human brain may be implemented via a network that includes bilateral DLPFC and bilateral IOS/IPS as key nodes.

Selecting and perceiving multiple visual objects

To explain how multiple visual objects are attended and perceived, we propose that our visual system first selects a fixed number of about four objects from a crowded scene based on their spatial information (object individuation) and then encode their details (object identification). We describe the involvement of the inferior intra-parietal sulcus (IPS) in object individuation and the superior IPS and higher visual areas in object identification. Our neural object-file theory synthesizes and extends existing ideas in visual cognition and is supported by behavioral and neuroimaging results. It provides a better understanding of the role of the different parietal areas in encoding visual objects and can explain various forms of capacity-limited processing in visual cognition such as working memory.

Distinctive Neural Mechanisms Supporting Visual Object Individuation and Identification

Many everyday activities, such as driving on a busy street, require the encoding of distinctive visual objects from crowded scenes. Given resource limitations of our visual system, one solution to this difficult and challenging task is to first select individual objects from a crowded scene (object individuation) and then encode their details (object identification). Using functional magnetic resonance imaging, two distinctive brain mechanisms were recently identified that support these two stages of visual object processing. While the inferior intraparietal sulcus (IPS) selects a fixed number of about four objects via their spatial locations, the superior IPS and the lateral occipital complex (LOC) encode the features of a subset of the selected objects in great detail (object shapes in this case). Thus, the inferior IPS individuates visual objects from a crowded display and the superior IPS and higher visual areas participate in subsequent object identification. Consistent with the prediction of this theory, even when only object shape identity but not its location is task relevant, this study shows that object individuation in the inferior IPS treats four identical objects similarly as four objects that are all different, whereas object shape identification in the superior IPS and the LOC treat four identical objects as a single unique object. These results provide independent confirmation supporting the dissociation between visual object individuation and identification in the brain.