Superior Intraparietal Sulcus Research Articles

Predicting Identity-Preserving Object Transformations in Human Posterior Parietal Cortex and Convolutional Neural Networks.

Previous research shows that, within human occipito-temporal cortex (OTC), we can use a general linear mapping function to link visual object responses across nonidentity feature changes, including Euclidean features (e.g., position and size) and non-Euclidean features (e.g., image statistics and spatial frequency). Although the learned mapping is capable of predicting responses of objects not included in training, these predictions are better for categories included than those not included in training. These findings demonstrate a near-orthogonal representation of object identity and nonidentity features throughout human OTC. Here, we extended these findings to examine the mapping across both Euclidean and non-Euclidean feature changes in human posterior parietal cortex (PPC), including functionally defined regions in inferior and superior intraparietal sulcus. We additionally examined responses in five convolutional neural networks (CNNs) pretrained with object classification, as CNNs are considered as the current best model of the primate ventral visual system. We separately compared results from PPC and CNNs with those of OTC. We found that a linear mapping function could successfully link object responses in different states of nonidentity transformations in human PPC and CNNs for both Euclidean and non-Euclidean features. Overall, we found that object identity and nonidentity features are represented in a near-orthogonal, rather than complete-orthogonal, manner in PPC and CNNs, just like they do in OTC. Meanwhile, some differences existed among OTC, PPC, and CNNs. These results demonstrate the similarities and differences in how visual object information across an identity-preserving image transformation may be represented in OTC, PPC, and CNNs.

The Human Intraparietal Sulcus Modulates Task-Evoked Functional Connectivity.

Past studies have demonstrated that flexible interactions between brain regions support a wide range of goal-directed behaviors. However, the neural mechanisms that underlie adaptive communication between brain regions are not well understood. In this study, we combined theta-burst transcranial magnetic stimulation (TMS) and functional magnetic resonance imaging to investigate the sources of top-down biasing signals that influence task-evoked functional connectivity. Subjects viewed sequences of images of faces and buildings and were required to detect repetitions (2-back vs. 1-back) of the attended stimuli category (faces or buildings). We found that functional connectivity between ventral temporal cortex and the primary visual cortex (VC) increased during processing of task-relevant stimuli, especially during higher memory loads. Furthermore, the strength of functional connectivity was greater for correct trials. Increases in task-evoked functional connectivity strength were correlated with increases in activity in multiple frontal, parietal, and subcortical (caudate and thalamus) regions. Finally, we found that TMS to superior intraparietal sulcus (IPS), but not to primary somatosensory cortex, decreased task-specific modulation in connectivity patterns between the primary VC and the parahippocampal place area. These findings demonstrate that the human IPS is a source of top-down biasing signals that modulate task-evoked functional connectivity among task-relevant cortical regions.

Task-context-dependent Linear Representation of Multiple Visual Objects in Human Parietal Cortex.

A host of recent studies have reported robust representations of visual object information in the human parietal cortex, similar to those found in ventral visual cortex. In ventral visual cortex, both monkey neurophysiology and human fMRI studies showed that the neural representation of a pair of unrelated objects can be approximated by the averaged neural representation of the constituent objects shown in isolation. In this study, we examined whether such a linear relationship between objects exists for object representations in the human parietal cortex. Using fMRI and multivoxel pattern analysis, we examined object representations in human inferior and superior intraparietal sulcus, two parietal regions previously implicated in visual object selection and encoding, respectively. We also examined responses from the lateral occipital region, a ventral object processing area. We obtained fMRI response patterns to object pairs and their constituent objects shown in isolation while participants viewed these objects and performed a 1-back repetition detection task. By measuring fMRI response pattern correlations, we found that all three brain regions contained representations for both single object and object pairs. In the lateral occipital region, the representation for a pair of objects could be reliably approximated by the average representation of its constituent objects shown in isolation, replicating previous findings in ventral visual cortex. Such a simple linear relationship, however, was not observed in either parietal region examined. Nevertheless, when we equated the amount of task information present by examining responses from two pairs of objects, we found that representations for the average of two object pairs were indistinguishable in both parietal regions from the average of another two object pairs containing the same four component objects but with a different pairing of the objects (i.e., the average of AB and CD vs. that of AD and CB). Thus, when task information was held consistent, the same linear relationship may govern how multiple independent objects are represented in the human parietal cortex as it does in ventral visual cortex. These findings show that object and task representations coexist in the human parietal cortex and characterize one significant difference of how visual information may be represented in ventral visual and parietal regions.

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.

Superior Intraparietal Sulcus Controls the Variability of Visual Working Memory Precision.

Limitations of working memory (WM) capacity depend strongly on the cognitive resources that are available for maintaining WM contents in an activated state. Increasing the number of items to be maintained in WM was shown to reduce the precision of WM and to increase the variability of WM precision over time. Although WM precision was recently associated with neural codes particularly in early sensory cortex, we have so far no understanding of the neural bases underlying the variability of WM precision, and how WM precision is preserved under high load. To fill this gap, we combined human fMRI with computational modeling of behavioral performance in a delayed color-estimation WM task. Behavioral results replicate a reduction of WM precision and an increase of precision variability under high loads (5 > 3 > 1 colors). Load-dependent BOLD signals in primary visual cortex (V1) and superior intraparietal sulcus (IPS), measured during the WM task at 2-4 s after sample onset, were modulated by individual differences in load-related changes in the variability of WM precision. Although stronger load-related BOLD increase in superior IPS was related to lower increases in precision variability, thus stabilizing WM performance, the reverse was observed for V1. Finally, the detrimental effect of load on behavioral precision and precision variability was accompanied by a load-related decline in the accuracy of decoding the memory stimuli (colors) from left superior IPS. We suggest that the superior IPS may contribute to stabilizing visual WM performance by reducing the variability of memory precision in the face of higher load. This study investigates the neural bases of capacity limitations in visual working memory by combining fMRI with cognitive modeling of behavioral performance, in human participants. It provides evidence that the superior intraparietal sulcus (IPS) is a critical brain region that influences the variability of visual working memory precision between and within individuals (Fougnie et al., 2012; van den Berg et al., 2012) under increased memory load, possibly in cooperation with perceptual systems of the occipital cortex. These findings substantially extend our understanding of the nature of capacity limitations in visual working memory and their neural bases. Our work underlines the importance of integrating cognitive modeling with univariate and multivariate methods in fMRI research, thus improving our knowledge of brain-behavior relationships.

Feature-Based Change Detection Reveals Inconsistent Individual Differences in Visual Working Memory Capacity.

Visual working memory (VWM) is a key cognitive system that enables people to hold visual information in mind after a stimulus has been removed and compare past and present to detect changes that have occurred. VWM is severely capacity limited to around 3–4 items, although there are robust individual differences in this limit. Importantly, these individual differences are evident in neural measures of VWM capacity. Here, we capitalized on recent work showing that capacity is lower for more complex stimulus dimension. In particular, we asked whether individual differences in capacity remain consistent if capacity is shifted by a more demanding task, and, further, whether the correspondence between behavioral and neural measures holds across a shift in VWM capacity. Participants completed a change detection (CD) task with simple colors and complex shapes in an fMRI experiment. As expected, capacity was significantly lower for the shape dimension. Moreover, there were robust individual differences in behavioral estimates of VWM capacity across dimensions. Similarly, participants with a stronger BOLD response for color also showed a strong neural response for shape within the lateral occipital cortex, intraparietal sulcus (IPS), and superior IPS. Although there were robust individual differences in the behavioral and neural measures, we found little evidence of systematic brain-behavior correlations across feature dimensions. This suggests that behavioral and neural measures of capacity provide different views onto the processes that underlie VWM and CD. Recent theoretical approaches that attempt to bridge between behavioral and neural measures are well positioned to address these findings in future work.

Decoding the content of visual short-term memory under distraction in occipital and parietal areas.

Recent studies have provided conflicting accounts regarding where in the human brain visual short-term memory (VSTM) content is stored, with strong univariate fMRI responses reported in superior intraparietal sulcus (IPS) but robust multivariate decoding reported in occipital cortex. Given the continuous influx of information in everyday vision, VSTM storage under distraction is often required. We found that neither distractor presence nor predictability during the memory delay affected behavioral performance. Similarly, superior IPS exhibited consistent decoding of VSTM content across all distractor manipulations and had multivariate responses that closely tracked behavioral VSTM performance. However, occipital decoding of VSTM content was significantly modulated by distractor presence and predictability. Furthermore, we found no effect of target-distractor similarity on VSTM behavioral performance, further challenging the role of sensory regions in VSTM storage. Overall, consistent with previous univariate findings, these results show that superior IPS, not occipital cortex, plays a central role in VSTM storage.

Optic Ataxia: From Balint’s Syndrome to the Parietal Reach Region

Optic ataxia is a high-order deficit in reaching to visual goals that occurs with posterior parietal cortex (PPC) lesions. It is a component of Balint's syndrome that also includes attentional and gaze disorders. Aspects of optic ataxia are misreaching in the contralesional visual field, difficulty preshaping the hand for grasping, and an inability to correct reaches online. Recent research in nonhuman primates (NHPs) suggests that many aspects of Balint's syndrome and optic ataxia are a result of damage to specific functional modules for reaching, saccades, grasp, attention, and state estimation. The deficits from large lesions in humans are probably composite effects from damage to combinations of these functional modules. Interactions between these modules, either within posterior parietal cortex or downstream within frontal cortex, may account for more complex behaviors such as hand-eye coordination and reach-to-grasp.

Gains of item-specific training in visual working memory and their neural correlates

Experts sometimes show higher working memory performance than novices but contrary to this finding, evidence for a positive effect of item-specific training is rare. This study provides evidence for item-specific training gains. We presented Chinese characters and artificial patterns (spotted figures) in a change detection task before and after training (varying set size from 1 to 3). A part of the Chinese characters were trained; others and the spotted figures were not trained. Memory capacity was between one and two items. For set size two, memory performance for trained characters was higher than for untrained characters and they were processed faster. Within superior intraparietal sulcus and middle occipital cortex (part of the putative posterior working memory network), the neural activity asymptotically increased with set size. Untrained items reached the activation maximum already at set size two. For this set size, the activity was significantly reduced for trained items so that a further increase from two to three items was observed. We interpret this difference as a correlate of a gain in neural efficiency. The size of this difference correlated with the training gain in memory. We assume that training causes a more efficient neural representation of trained items supported by long-term memory and this allows holding more items in working memory.

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 search for the neural mechanisms of the set size effect

The set size effect in visual search refers to the linear increase in response time (RT) or decrease in accuracy as the number of distractors increases. Previous human and monkey studies have reported a correlation between set size and neural activity in the frontal eye field (FEF) and intraparietal sulcus (IPS). In a recent functional magnetic resonance imaging study, we did not observe a set size effect in the superior precentral sulcus (sPCS, thought to be the human homolog of the FEF) and IPS in an oculomotor visual search task (Ikkai et al., 2011). Our task used placeholders in the search array, along with the target and distractors, in order to equate the amount of retinal stimulation for each set size. We here attempted to reconcile these differences with the results from a follow-up experiment in which the same oculomotor visual search task was used, but without placeholders. A strong behavioral set size effect was observed in both studies, with very similar saccadic RTs and slopes between RT and set size. However, a set size effect was now observed in the sPCS and IPS. We comment on this finding and discuss the role of these neural areas in visual search.

Functional MRI evidence for the importance of visual short‐term memory in logographic reading

Logographic symbols are visually complex, and thus children's abilities for visual short-term memory (VSTM) predict their reading competence in logographic systems. In the present study, we investigated the importance of VSTM in logographic reading in adults, both behaviorally and by means of fMRI. Outside the scanner, VSTM predicted logographic Kanji reading in native Japanese adults (n=45), a finding consistent with previous observations in Japanese children. In the scanner, participants (n=15) were asked to perform a visual one-back task. For this fMRI experiment, we took advantage of the unique linguistic characteristic of the Japanese writing system, whereby syllabic Kana and logographic Kanji can share the same sound and meaning, but differ only in the complexity of their visual features. Kanji elicited greater activation than Kana in the cerebellum and two regions associated with VSTM, the lateral occipital complex and the superior intraparietal sulcus, bilaterally. The same regions elicited the highest activation during the control condition (an unfamiliar, unpronounceable script to the participants), presumably due to the increased VSTM demands for processing the control script. In addition, individual differences in VSTM performance (outside the scanner) significantly predicted blood oxygen level-dependent signal changes in the identified VSTM regions, during the Kanji and control conditions, but not during the Kana condition. VSTM appears to play an important role in reading logographic words, even in skilled adults, as evidenced at the behavioral and neural level, most likely due to the increased VSTM/visual attention demands necessary for processing complex visual features inherent in logographic symbols.

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.