Human Intraparietal Sulcus Research Articles

Systems-level decoding reveals the cognitive and behavioral profile of the human intraparietal sulcus.

The human intraparietal sulcus (IPS) covers large portions of the posterior cortical surface and has been implicated in a variety of cognitive functions. It is, however, unclear how cognitive functions dissociate between the IPS's heterogeneous subdivisions, particularly in perspective to their connectivity profile. We applied a neuroinformatics driven system-level decoding on three cytoarchitectural distinct subdivisions (hIP1, hIP2, hIP3) per hemisphere, with the aim to disentangle the cognitive profile of the IPS in conjunction with functionally connected cortical regions. The system-level decoding revealed nine functional systems based on meta-analytical associations of IPS subdivisions and their cortical coactivations: Two systems-working memory and numeric cognition-which are centered on all IPS subdivisions, and seven systems-attention, language, grasping, recognition memory, rotation, detection of motions/shapes and navigation-with varying degrees of dissociation across subdivisions and hemispheres. By probing the spatial overlap between systems-level co-activations of the IPS and seven canonical intrinsic resting state networks, we observed a trend toward more co-activation between hIP1 and the front parietal network, between hIP2 and hIP3 and the dorsal attention network, and between hIP3 and the visual and somatomotor network. Our results confirm previous findings on the IPS's role in cognition but also point to previously unknown differentiation along the IPS, which present viable starting points for future work. We also present the systems-level decoding as promising approach toward functional decoding of the human connectome.

Connectivity-Defined Subdivisions of the Intraparietal Sulcus Respond Differentially to Abstraction during Decision-Making.

The intraparietal sulcus (IPS) has been implicated in numerous functions that range from representation of visual stimuli to action planning, but its role in abstract decision-making has been unclear, in part because low-level functions often act as confounds. Here, we address this problem using a task that dissociates abstract decision-making from sensory salience, attentional control, motor planning, and motor output. Functional MRI data were collected from healthy female and male human subjects while they performed a policy abstraction task requiring use of a more abstract (second-order) rule to select between two less abstract (first-order) rules that informed the motor response. By identifying IPS subdivisions with preferential connectivity to prefrontal regions that are differentially responsive to task abstraction, we found that a caudal IPS (cIPS) subregion with strongest connectivity to the pre-premotor cortex was preferentially active for second-order cues, whereas a rostral IPS subregion (rIPS) with strongest connectivity to the dorsal premotor cortex was active during attentional control over first-order cues. These effects for abstraction were seen in addition to cIPS activity that was specific to sensory salience, and rIPS activity that was specific to motor output. Notably, topographic responses to the second-order cue were detected along the caudal-rostral axis of IPS, mirroring the broader organization seen in lateral prefrontal cortex. Together, these data demonstrate that subregions within IPS exhibit activity responsive to policy abstraction, and they suggest that IPS may be organized into frontoparietal subnetworks that support hierarchical cognitive control.SIGNIFICANCE STATEMENT Abstract decision-making allows us to flexibly adapt our behavior to new contexts. Although much previous work has focused on the role of lateral prefrontal cortex in such decisions, the contributions of parietal cortex have been relatively understudied. Here, we demonstrate that spatially segregated subregions of human IPS with strong functional connections to lateral prefrontal cortex demonstrate activity selective for abstract decisions. This activity can be distinguished from responses resulting from cognitive processes related to sensory salience, attentional control, motor planning, and movement. Together, these findings indicate that different task demands are reflected in the topography of IPS, and they explicitly reveal a role in abstract decision-making.

Representations for grasp-relevant parts of objects in the human intraparietal sulcus

Representations for grasp-relevant parts of objects in the human intraparietal sulcus

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.

Individual retinotopic organization in human intraparietal sulcus predicted by connectivity fingerprinting

Individual retinotopic organization in human intraparietal sulcus predicted by connectivity fingerprinting

Chapter 3 - Microarchitecture and connectivity of the parietal lobe

This chapter summarizes current knowledge on the structural segregation of the parietal lobe based on cyto-, myelo-, and receptorarchitectonic studies, as well as the connectivity of this brain region with other cortical and subcortical structures. The anterior part of the human parietal cortex comprises the somatosensory areas 3a, 3b, 1, and 2, whereas the posterior part contains seven multimodal areas in both the superior and inferior parietal lobules. Available cytoarchitectonic maps of the human intraparietal sulcus do not provide a complete picture yet. Myelo- and receptorarchitectonic analyses largely confirm but also further differentiate the cytoarchitectonic maps. With the advent of diffusion imaging and functional connectivity studies, further insight into the structural and functional organization has been achieved. It shows that the posterior parietal cortex is a key node in anatomic networks connecting visual with (pre)frontal cortices, and temporal with parts of frontal cortices. Here, the superior longitudinal fascicle and its components play a major role, together with the arcuate and middle longitudinal fascicles. Major connections with subcortical structures, particularly the basal ganglia and thalamic nuclei, are discussed. Finally, the importance of precise maps of parietal areas for defining seed regions in structural and functional connectivity studies is emphasized.

Functional subdivisions within the human intraparietal sulcus are involved in visuospatial transformation in a non-context-dependent manner.

Object-based visuospatial transformation is important for the ability to interact with the world and the people and objects within it. In this preliminary investigation, we hypothesized that object-based visuospatial transformation is a unitary process invoked regardless of current context and is localized to the intraparietal sulcus. Participants (n = 14) performed both antisaccade and mental rotation tasks while scanned using fMRI. A statistical conjunction confirmed that both tasks activated the intraparietal sulcus. Statistical parametric anatomical mapping determined that the statistical conjunction was localized to intraparietal sulcus subregions hIP2 and hIP3. A Gaussian naïve Bayes classifier confirmed that the conjunction in region hIP3 was indistinguishable between tasks. The results provide evidence that object-based visuospatial transformation is a domain-general process that is invoked regardless of current context. Our results are consistent with the modular model of the posterior parietal cortex and the distinct cytoarchitectonic, structural, and functional connectivity profiles of the subregions in the intraparietal sulcus. Hum Brain Mapp 39:354-368, 2018. © 2017 Wiley Periodicals, Inc.

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.

Probing the reaching-grasping network in humans through multivoxel pattern decoding.

The quest for a putative human homolog of the reaching-grasping network identified in monkeys has been the focus of many neuropsychological and neuroimaging studies in recent years. These studies have shown that the network underlying reaching-only and reach-to-grasp movements includes the superior parieto-occipital cortex (SPOC), the anterior part of the human intraparietal sulcus (hAIP), the ventral and the dorsal portion of the premotor cortex, and the primary motor cortex (M1). Recent evidence for a wider frontoparietal network coding for different aspects of reaching-only and reach-to-grasp actions calls for a more fine-grained assessment of the reaching-grasping network in humans by exploiting pattern decoding methods (multivoxel pattern analysis--MVPA). Here, we used MPVA on functional magnetic resonance imaging (fMRI) data to assess whether regions of the frontoparietal network discriminate between reaching-only and reach-to-grasp actions, natural and constrained grasping, different grasp types, and object sizes. Participants were required to perform either reaching-only movements or two reach-to-grasp types (precision or whole hand grasp) upon spherical objects of different sizes. Multivoxel pattern analysis highlighted that, independently from the object size, all the selected regions of both hemispheres contribute in coding for grasp type, with the exception of SPOC and the right hAIP. Consistent with recent neurophysiological findings on monkeys, there was no evidence for a clear-cut distinction between a dorsomedial and a dorsolateral pathway that would be specialized for reaching-only and reach-to-grasp actions, respectively. Nevertheless, the comparison of decoding accuracy across brain areas highlighted their different contributions to reaching-only and grasping actions. Altogether, our findings enrich the current knowledge regarding the functional role of key brain areas involved in the cortical control of reaching-only and reach-to-grasp actions in humans, by revealing novel fine-grained distinctions among action types within a wide frontoparietal network.

Influences of Long-Term Memory-Guided Attention and Stimulus-Guided Attention on Visuospatial Representations within Human Intraparietal Sulcus.

The human parietal lobe contains multiple maps of the external world that spatially guide perception, action, and cognition. Maps in each cerebral hemisphere code information from the opposite side of space, not from the same side, and the two hemispheres are symmetric. Paradoxically, damage to specific parietal regions that lack spatial maps can cause patients to ignore half of space (hemispatial neglect syndrome), but only for right (not left) hemisphere damage. Conversely, the left parietal cortex has been linked to retrieval of vivid memories regardless of space. Here, we investigate possible underlying mechanisms in healthy individuals. We demonstrate two forms of dynamic changes in parietal spatial representations: an asymmetric one for stimulus-guided attention and a symmetric one for long-term memory-guided attention.

Visual number beats abstract numerical magnitude: format-dependent representation of Arabic digits and dot patterns in human parietal cortex.

In numerical cognition, there is a well-known but contested hypothesis that proposes an abstract representation of numerical magnitude in human intraparietal sulcus (IPS). On the other hand, researchers of object cognition have suggested another hypothesis for brain activity in IPS during the processing of number, namely that this activity simply correlates with the number of visual objects or units that are perceived. We contrasted these two accounts by analyzing multivoxel activity patterns elicited by dot patterns and Arabic digits of different magnitudes while participants were explicitly processing the represented numerical magnitude. The activity pattern elicited by the digit "8" was more similar to the activity pattern elicited by one dot (with which the digit shares the number of visual units but not the magnitude) compared to the activity pattern elicited by eight dots, with which the digit shares the represented abstract numerical magnitude. A multivoxel pattern classifier trained to differentiate one dot from eight dots classified all Arabic digits in the one-dot pattern category, irrespective of the numerical magnitude symbolized by the digit. These results were consistently obtained for different digits in IPS, its subregions, and many other brain regions. As predicted from object cognition theories, the number of presented visual units forms the link between the parietal activation elicited by symbolic and nonsymbolic numbers. The current study is difficult to reconcile with the hypothesis that parietal activation elicited by numbers would reflect a format-independent representation of number.

Early intraparietal involvement in motion-driven attention identified with fMRI-neuronavigated TMS

Event Abstract Back to Event Early intraparietal involvement in motion-driven attention identified with fMRI-neuronavigated TMS Bonnie Alexander1, Robin Laycock1, Sheila Crewther1* and David Crewther2 1 La Trobe University, Australia 2 Swinburne University, Australia Background Human intraparietal sulcus (IPS) is a key region involved in visual attention. In investigations of the timing of visual processing, both early and late critical periods of activity post-stimulus have been identified in multiple cortical areas (e.g. Corthout, Uttl, Ziemann, Cowey, & Hallett, 1998; Ioannides & Poghosyan, 2012; Morand et al., 2000) including area V5 for motion-based stimuli identified using chronometric TMS (Laycock, Crewther, Fitzgerald, & Crewther, 2007). The current study aimed to determine whether similar critical periods could be detected using fMRI-neuronavigated chronometric TMS applied to areas of IPS. Methods Three women and six men (age M = 28.5, SD = 3.38) participated in fMRI and TMS testing. fMRI was utilised to identify V5, and middle and posterior intraparietal sulcus (mIPS; pIPS) regions active during a motion-driven attention task. These areas were defined individually per participant and used for TMS neuronavigation. Paired-pulse TMS was applied during task performance at 30 ms stimulus onset asynchronies (SOAs) between 0-180 ms. Relative decreases in accuracy were used to indicate critical times of involvement of each area. Results For TMS applied to mIPS, there was a relative decrease in performance at the 150 ms SOA, trends towards decreases at 30 and 60 ms, as well as a relative increase at 180 ms. TMS to pIPS revealed a relative decrease in performance at 0 ms and a trend towards a drop at 30 ms. There were no significant decreases in performance for TMS to V5 at any SOA. Conclusions The results for mIPS TMS are consistent with findings of multiple phases of activity following stimulus presentation, i.e. a possible early (0-30 ms) and late (150ms) phase. The early decreases seen for TMS to pIPS could be interpreted in terms of prestimulus anticipatory attention processes. These results are consistent with early timepoints of activity found in other studies, and, if replicable, could be incorporated into timing models of visual function. Keywords: Intraparietal sulcus, chronometric TMS, human visual system, fMRI-neuronavigated TMS, motion-driven attention Conference: XII International Conference on Cognitive Neuroscience (ICON-XII), Brisbane, Queensland, Australia, 27 Jul - 31 Jul, 2014. Presentation Type: Poster Topic: Attention Citation: Alexander B, Laycock R, Crewther S and Crewther D (2015). Early intraparietal involvement in motion-driven attention identified with fMRI-neuronavigated TMS. Conference Abstract: XII International Conference on Cognitive Neuroscience (ICON-XII). doi: 10.3389/conf.fnhum.2015.217.00077 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 19 Feb 2015; Published Online: 24 Apr 2015. * Correspondence: Prof. Sheila Crewther, La Trobe University, Melbourne, Australia, s.crewther@latrobe.edu.au Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Bonnie Alexander Robin Laycock Sheila Crewther David Crewther Google Bonnie Alexander Robin Laycock Sheila Crewther David Crewther Google Scholar Bonnie Alexander Robin Laycock Sheila Crewther David Crewther PubMed Bonnie Alexander Robin Laycock Sheila Crewther David Crewther Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Global versus local: double dissociation between MT+ and V3A in motion processing revealed using continuous theta burst transcranial magnetic stimulation.

The functional properties of motion selective areas in human visual cortex, including V3A, MT+, and intraparietal sulcus (IPS) are not fully understood. To examine the functional specialization of these areas for global and local motion processing, we used off-line, neuronavigated, continuous theta burst (cTBS) transcranial magnetic stimulation to temporarily alter neural activity within unilateral V3A, MT+, and IPS. A within-subjects design was employed and stimulation sessions were separated by at least 24 h. In each session, subjects were asked to discriminate the global motion directions of successively presented random dot kinematograms (RDKs) before and after cTBS. RDKs were presented at either 100 or 40 % coherence in either the left or right visual field. We found that V3A stimulation selectively impaired discrimination of 100 % coherent motion, while MT+ stimulation selectively impaired discrimination of 40 % coherent motion. IPS stimulation impaired discrimination of both motion stimuli. All cTBS effects were specific to stimuli presented contralaterally to the stimulation site and vertex stimulation had no effect. The double dissociation between the cTBS effects on MT+ and V3A indicates distinct roles for these two regions in motion processing. Judging the direction of 100 % coherent motion can rely on local motion processing because every dot moves in the same direction. However, judging the global direction of 40 % coherent motion requires global processing. Thus, our results suggest separate, parallel processing of local and global motion in V3A and MT+, respectively, with the outputs of these two areas being combined within the IPS.

Differential roles of the frontal and parietal cortices in the control of saccades

Although externally as well as internally-guided eye movements allow us to flexibly explore the visual environment, their differential neural mechanisms remain elusive. A better understanding of these neural mechanisms will help us to understand the control of action and to elucidate the nature of cognitive deficits in certain psychiatric populations (e.g. schizophrenia) that show increased latencies in volitional but not visually-guided saccades. Both the superior precentral sulcus (sPCS) and the intraparietal sulcus (IPS) are implicated in the control of eye movements. However, it remains unknown what differential contributions the two areas make to the programming of visually-guided and internally-guided saccades. In this study we tested the hypotheses that sPCS and IPS distinctly encode internally-guided saccades and visually-guided saccades. We scanned subjects with fMRI while they generated visually-guided and internally-guided delayed saccades. We used multi-voxel pattern analysis to test whether patterns of cue related, preparatory and saccade related activation could be used to predict the direction of the planned eye movement. Results indicate that patterns in the human sPCS predicted internally-guided saccades but not visually-guided saccades in all trial periods and patterns in the IPS predicted internally-guided saccades and visually-guided saccades equally well. The results support the hypothesis that the human sPCS and IPS make distinct contributions to the control of volitional eye movements.

The representation of tool and non-tool object information in the human intraparietal sulcus

Humans have an amazing ability to quickly and efficiently recognize and interact with visual objects in their environment. The underlying neural processes supporting this ability have been mainly explored in the ventral visual stream. However, the dorsal stream has been proposed to play a critical role in guiding object-directed actions. This hypothesis is supported by recent neuroimaging studies that have identified object-selective and tool-related activity in human parietal cortex. In the present study, we sought to delineate tool-related information in the anterior portions of the human intraparietal sulcus (IPS) and relate it to recently identified motor-defined and topographic regions of interest (ROIs) using functional MRI in individual subjects. Consistent with previous reports, viewing pictures of tools compared with pictures of animals led to a higher blood oxygenation level-dependent (BOLD) response in the left anterior IPS. For every subject, this activation was located lateral, anterior, and inferior to topographic area IPS5 and lateral and inferior to a motor-defined human parietal grasp region (hPGR). In a separate experiment, subjects viewed pictures of tools, animals, graspable (non-tool) objects, and scrambled objects. An ROI-based time-course analysis showed that tools evoked a stronger BOLD response than animals throughout topographic regions of the left IPS. Additionally, graspable objects evoked stronger responses than animals, equal to responses to tools, in posterior regions and weaker responses than tools, equal to responses to animals, in anterior regions. Thus the left anterior tool-specific region may integrate visual information encoding graspable features of objects from more posterior portions of the IPS with experiential knowledge of object use and function to guide actions.

Functional organization of human posterior parietal cortex: grasping- and reaching-related activations relative to topographically organized cortex

The act of reaching to grasp an object requires the coordination between transporting the arm and shaping the hand. Neurophysiological, neuroimaging, neuroanatomic, and neuropsychological studies in macaque monkeys and humans suggest that the neural networks underlying grasping and reaching acts are at least partially separable within the posterior parietal cortex (PPC). To better understand how these neural networks have evolved in primates, we characterized the relationship between grasping- and reaching-related responses and topographically organized areas of the human intraparietal sulcus (IPS) using functional MRI. Grasping-specific activation was localized to the left anterior IPS, partially overlapping with the most anterior topographic regions and extending into the postcentral sulcus. Reaching-specific activation was localized to the left precuneus and superior parietal lobule, partially overlapping with the medial aspects of the more posterior topographic regions. Although the majority of activity within the topographic regions of the IPS was nonspecific with respect to movement type, we found evidence for a functional gradient of specificity for reaching and grasping movements spanning posterior-medial to anterior-lateral PPC. In contrast to the macaque monkey, grasp- and reach-specific activations were largely located outside of the human IPS.

Common parietal activation in musical mental transformations across pitch and time

We previously observed that mental manipulation of the pitch level or temporal organization of melodies results in functional activation in the human intraparietal sulcus (IPS), a region also associated with visuospatial transformation and numerical calculation. Two outstanding questions about these musical transformations are whether pitch and time depend on separate or common processing in IPS, and whether IPS recruitment in melodic tasks varies depending upon the degree of transformation required (as it does in mental rotation). In the present study we sought to answer these questions by applying functional magnetic resonance imaging while musicians performed closely matched mental transposition (pitch transformation) and melody reversal (temporal transformation) tasks. A voxel-wise conjunction analysis showed that in individual subjects, both tasks activated overlapping regions in bilateral IPS, suggesting that a common neural substrate subserves both types of mental transformation. Varying the magnitude of mental pitch transposition resulted in variation of IPS BOLD signal in correlation with the musical key-distance of the transposition, but not with the pitch distance, indicating that the cognitive metric relevant for this type of operation is an abstract one, well described by music-theoretic concepts. These findings support a general role for the IPS in systematically transforming auditory stimulus representations in a nonspatial context.

Neural correlates of changing intention in the human FEF and IPS

Previous research demonstrates that our apparent mental flexibility depends largely on the strength of our prior intention; changing our intention in advance enables a smooth transition from one task to another (e.g., Astle DE, Jackson GM, Swainson R. J Cogn Neurosci 20: 255-267, 2008; Duncan J, Emslie H, Williams P, Johnson R, Freer C. Cogn Psychol 30: 257-303, 1996; Husain M, Parton A, Hodgson TL, Mort D, Rees G. Nat Neurosci 6: 117-118, 2003). However, these necessarily rapid anticipatory mechanisms have been difficult to study in the human brain. We used EEG and magnetoencephalography, specifically event-related potentials and fields (ERPs and ERFs), respectively, to explore the neural correlates of this important aspect of mental flexibility. Subjects performed a manual version of a pro/antisaccade task using preparatory cues to switch between the pro- and antirules. When subjects switched their intention, we observed a positivity over central electrodes, which correlated significantly with our behavioral data; the greater the ERP effect, the stronger the subject's change of intention. ERFs, alongside subject-specific structural MRIs, were used to project into source space. When subjects switched their intention, they showed significantly elevated activity in the right frontal eye field and left intraparietal sulcus (IPS); the greater the left IPS activity on switch trials, the stronger the subject's change of intention. This network has previously been implicated in the top-down control of eye movements, but here we demonstrate its role in the top-down control of a task set, in particular, that it is recruited when we change the task that we intend to perform.

The Benefit of Object Interactions Arises in the Lateral Occipital Cortex Independent of Attentional Modulation from the Intraparietal Sulcus: A Transcranial Magnetic Stimulation Study

Our visual experience is generally not of isolated objects, but of scenes, where multiple objects are interacting. Such interactions (e.g., a watering can positioned to pour water toward a plant) have been shown to facilitate object identification compared with when the objects are depicted as not interacting (e.g., a watering can positioned away from the plant) (Green and Hummel, 2004, 2006). What is the neural basis for this advantage? Recent fMRI studies have identified the lateral occipital cortex (LO) as a potential neural origin of this behavioral benefit, as LO showed greater responses to object pairs depicted as interacting compared with when they are not (Kim and Biederman, 2010; Roberts and Humphreys, 2010). However, it is possible that LO was modulated by an attention-sensitive region, the intraparietal sulcus (IPS), which sometimes showed a similar pattern of responses as that of LO in the Kim and Biederman (2010) investigation. To test this hypothesis, we delivered transcranial magnetic stimulation (TMS) to human subjects' LO and IPS while they detected a target object that was or was not interacting with another object to form a scene. TMS delivered to LO but not IPS abolished the facilitation in identifying interacting objects compared with noninteracting depictions observed in the absence of TMS, suggesting that it is LO and not IPS that is critical for the coding of object interactions.

Distinct representations of numerical and non-numerical order in the human intraparietal sulcus revealed by multivariate pattern recognition

Neuroimaging studies of numerical cognition have pointed to the horizontal segment of the intraparietal sulcus (hIPS) as the neural correlate of numerical representations in humans. However, the specificity of hIPS for numbers remains controversial. For example, its activation during numerical comparison cannot be distinguished from activation during ordinal judgments on non-numerical sequences such as letters (Fias et al., 2007, J. Neuroscience). Based on the hypothesis that the fine-grained distinction between representations of numerical vs. letter order in hIPS might simply be invisible to conventional fMRI data analysis, we used support vector machines (SVM) to reanalyse the data of Fias et al. (2007). We show that classifiers trained on hIPS voxels can discriminate between number comparison and letter comparison, even though the two tasks produce the same metric of behaviour. Voxels discriminating between the two conditions were consistent across subjects and contribution analysis revealed maps of distinct sets of voxels implicated in the processing of numerical vs. alphabetical order in bilateral hIPS. These results reconcile the neuroimaging data with the neuropsychological evidence suggesting dissociations between numbers and other non-numerical ordered sequences, and demonstrate that multivariate analyses are fundamental to address fine-grained theoretical issues with fMRI studies.