A supramodal number representation in human intraparietal cortex

The triple-code theory of numerical processing postulates an abstract-semantic "number sense." Neuropsychology points to intraparietal cortex as a potential substrate, but previous functional neuroimaging studies did not dissociate the representation of numerical magnitude from task-driven effects on intraparietal activation. In an event-related fMRI study, we presented numbers, letters, and colors in the visual and auditory modality, asking subjects to respond to target items within each category. In the absence of explicit magnitude processing, numbers compared with letters and colors across modalities activated a bilateral region in the horizontal intraparietal sulcus. This stimulus-driven number-specific intraparietal response supports the idea of a supramodal number representation that is automatically accessed by presentation of numbers and may code magnitude information.     

Impaired parietal magnitude processing in developmental dyscalculia

Developmental dyscalculia (DD) is a specific learning disability affecting the acquisition of school-level mathematical abilities in the context of otherwise normal academic achievement, with prevalence estimates in the order of 3-6%. Behavioural studies show deficits in elementary numerical processing among individuals with pure DD, indicating that deficits in higher-level mathematical skills may stem from impaired representation and processing of basic numerical magnitude. Adult neuropsychological and neuroimaging research points to the intraparietal sulcus as a key region for the representation and processing of numerical magnitude. This raises the possibility of a parietal dysfunction as a root cause of DD. We show that, in children with pure DD, the right intraparietal sulcus is not modulated in response to numerical processing demands to the same degree as in typically developing children. This finding provides the first direct evidence for a specific impairment of parietal magnitude systems in DD during non-symbolic numerosity processing.     

Does the Parietal Cortex Distinguish between “10,” “Ten,” and Ten Dots?

It is well established that the intraparietal sulcus (IPS) plays an important role in the processing and representation of numerical magnitude. Two recent studies by Piazza et al. and Cohen Kadosh et al. published in this issue of Neuron used fMRI adaptation to explore the extent to which parietal number processing is dependent upon or independent of a numbers' surface format. Their results, while slightly different, converge to suggest that the answer may be neither, but rather that it depends on the hemisphere.     

Neural basis of genetically determined visuospatial construction deficit in Williams syndrome

A unique opportunity to understand genetic determinants of cognition is offered by Williams syndrome (WS), a well-characterized hemideletion on chromosome 7q11.23 that causes extreme, specific weakness in visuospatial construction (the ability to visualize an object as a set of parts or construct a replica). Using multimodal neuroimaging, we identified a neural mechanism underlying the WS visuoconstructive deficit. Hierarchical assessment of visual processing with fMRI showed isolated hypoactivation in WS in the parietal portion of the dorsal stream. In the immediately adjacent parietooccipital/intraparietal sulcus, structural neuroimaging showed a gray matter volume reduction in participants with WS. Path analysis demonstrated that the functional abnormalities could be attributed to impaired input from this structurally altered region. Our observations confirm a longstanding hypothesis about dorsal stream dysfunction in WS, demonstrate effects of a localized abnormality on visual information processing in humans, and define a systems-level phenotype for mapping genetic determinants of visuoconstructive function.     

Uncovering the spatio-temporal dynamics of value-based decision-making in the human brain: a combined fMRI–EEG study

While there is a growing body of functional magnetic resonance imaging (fMRI) evidence implicating a corpus of brain regions in value-based decision-making in humans, the limited temporal resolution of fMRI cannot address the relative temporal precedence of different brain regions in decision-making. To address this question, we adopted a computational model-based approach to electroencephalography (EEG) data acquired during a simple binary choice task. fMRI data were also acquired from the same participants for source localization. Post-decision value signals emerged 200 ms post-stimulus in a predominantly posterior source in the vicinity of the intraparietal sulcus and posterior temporal lobe cortex, alongside a weaker anterior locus. The signal then shifted to a predominantly anterior locus 850 ms following the trial onset, localized to the ventromedial prefrontal cortex and lateral prefrontal cortex. Comparison signals between unchosen and chosen options emerged late in the trial at 1050 ms in dorsomedial prefrontal cortex, suggesting that such comparison signals may not be directly associated with the decision itself but rather may play a role in post-decision action selection. Taken together, these results provide us new insights into the temporal dynamics of decision-making in the brain, suggesting that for a simple binary choice task, decisions may be encoded predominantly in posterior areas such as intraparietal sulcus, before shifting anteriorly.     

Spatio-Temporal Dynamics of Human Intention Understanding in Temporo-Parietal Cortex: A Combined EEG/fMRI Repetition Suppression Paradigm

Inferring the intentions of other people from their actions recruits an inferior fronto-parietal action observation network as well as a putative social network that includes the posterior superior temporal sulcus (STS). However, the functional dynamics within and among these networks remains unclear. Here we used functional magnetic resonance imaging (fMRI) and high-density electroencephalogram (EEG), with a repetition suppression design, to assess the spatio-temporal dynamics of decoding intentions. Suppression of fMRI activity to the repetition of the same intention was observed in inferior frontal lobe, anterior intraparietal sulcus (aIPS), and right STS. EEG global field power was reduced with repeated intentions at an early (starting at 60 ms) and a later (∼330 ms) period after the onset of a hand-on-object encounter. Source localization during these two intervals involved right STS and aIPS regions highly consistent with RS effects observed with fMRI. These results reveal the dynamic involvement of temporal and parietal networks at multiple stages during the intention decoding and without a strict segregation of intention decoding between these networks.     

The neural correlates of problem states: testing FMRI predictions of a computational model of multitasking

Background

It has been shown that people can only maintain one problem state, or intermediate mental representation, at a time. When more than one problem state is required, for example in multitasking, performance decreases considerably. This effect has been explained in terms of a problem state bottleneck.

Methodology

In the current study we use the complimentary methodologies of computational cognitive modeling and neuroimaging to investigate the neural correlates of this problem state bottleneck. In particular, an existing computational cognitive model was used to generate a priori fMRI predictions for a multitasking experiment in which the problem state bottleneck plays a major role. Hemodynamic responses were predicted for five brain regions, corresponding to five cognitive resources in the model. Most importantly, we predicted the intraparietal sulcus to show a strong effect of the problem state manipulations.

Conclusions

Some of the predictions were confirmed by a subsequent fMRI experiment, while others were not matched by the data. The experiment supported the hypothesis that the problem state bottleneck is a plausible cause of the interference in the experiment and that it could be located in the intraparietal sulcus.     

Functional imaging of numerical processing in adults and 4-y-old children

Adult humans, infants, pre-school children, and non-human animals appear to share a system of approximate numerical processing for non-symbolic stimuli such as arrays of dots or sequences of tones. Behavioral studies of adult humans implicate a link between these non-symbolic numerical abilities and symbolic numerical processing (e.g., similar distance effects in accuracy and reaction-time for arrays of dots and Arabic numerals). However, neuroimaging studies have remained inconclusive on the neural basis of this link. The intraparietal sulcus (IPS) is known to respond selectively to symbolic numerical stimuli such as Arabic numerals. Recent studies, however, have arrived at conflicting conclusions regarding the role of the IPS in processing non-symbolic, numerosity arrays in adulthood, and very little is known about the brain basis of numerical processing early in development. Addressing the question of whether there is an early-developing neural basis for abstract numerical processing is essential for understanding the cognitive origins of our uniquely human capacity for math and science. Using functional magnetic resonance imaging (fMRI) at 4-Tesla and an event-related fMRI adaptation paradigm, we found that adults showed a greater IPS response to visual arrays that deviated from standard stimuli in their number of elements, than to stimuli that deviated in local element shape. These results support previous claims that there is a neurophysiological link between non-symbolic and symbolic numerical processing in adulthood. In parallel, we tested 4-y-old children with the same fMRI adaptation paradigm as adults to determine whether the neural locus of non-symbolic numerical activity in adults shows continuity in function over development. We found that the IPS responded to numerical deviants similarly in 4-y-old children and adults. To our knowledge, this is the first evidence that the neural locus of adult numerical cognition takes form early in development, prior to sophisticated symbolic numerical experience. More broadly, this is also, to our knowledge, the first cognitive fMRI study to test healthy children as young as 4 y, providing new insights into the neurophysiology of human cognitive development.     

Human cortical regions involved in extracting depth from motion

We used functional magnetic resonance imaging (fMRI) to investigate brain regions involved in extracting three-dimensional structure from motion. A factorial design included two-dimensional and three-dimensional structures undergoing rigid and nonrigid motions. As predicted from monkey data, the human homolog of MT/V5 was significantly more active when subjects viewed three-dimensional (as opposed to two-dimensional) displays, irrespective of their rigidity. Human MT/V5+ (hMT/V5+) is part of a network with right hemisphere dominance involved in extracting depth from motion, including a lateral occipital region, five sites along the intraparietal sulcus (IPS), and two ventral occipital regions. Control experiments confirmed that this pattern of activation is most strongly correlated with perceived three-dimensional structure, in as much as it arises from motion and cannot be attributed to numerous two-dimensional image properties or to saliency.     

Maps of space in human frontoparietal cortex

Prefrontal cortex (PFC) and posterior parietal cortex (PPC) are neural substrates for spatial cognition. We here review studies in which we tested the hypothesis that human frontoparietal cortex may function as a priority map. According to priority map theory, objects or locations in the visual world are represented by neural activity that is proportional to their attentional priority. Using functional magnetic resonance imaging (fMRI), we first identified topographic maps in PFC and PPC as candidate priority maps of space. We then measured fMRI activity in candidate priority maps during the delay periods of a covert attention task, a spatial working memory task, and a motor planning task to test whether the activity depended on the particular spatial cognition. Our hypothesis was that some, but not all, candidate priority maps in PFC and PPC would be agnostic with regard to what was being prioritized, in that their activity would reflect the location in space across tasks rather than a particular kind of spatial cognition (e.g., covert attention). To test whether patterns of delay period activity were interchangeable during the spatial cognitive tasks, we used multivariate classifiers. We found that decoders trained to predict the locations on one task (e.g., working memory) cross-predicted the locations on the other tasks (e.g., covert attention and motor planning) in superior precentral sulcus (sPCS) and in a region of intraparietal sulcus (IPS2), suggesting that these patterns of maintenance activity may be interchangeable across the tasks. Such properties make sPCS in frontal cortex and IPS2 in parietal cortex viable priority map candidates, and suggest that these areas may be the human homologs of the monkey frontal eye field (FEF) and lateral intraparietal area (LIP).     

A magnitude code common to numerosities and number symbols in human intraparietal cortex

Activation of the horizontal segment of the intraparietal sulcus (hIPS) has been observed in various number-processing tasks, whether numbers were conveyed by symbolic numerals (digits, number words) or by nonsymbolic displays (dot patterns). This suggests an abstract coding of numerical magnitude. Here, we critically tested this hypothesis using fMRI adaptation to demonstrate notation-independent coding of numerical quantity in the hIPS. Once subjects were adapted either to dot patterns or to Arabic digits, activation in the hIPS and in frontal regions recovered in a distance-dependent fashion whenever a new number was presented, irrespective of notation changes. This remained unchanged when analyzing the hIPS peaks from an independent localizer scan of mental calculation. These results suggest an abstract coding of approximate number common to dots, digits, and number words. They support the idea that symbols acquire meaning by linking neural populations coding symbol shapes to those holding nonsymbolic representations of quantities.     

Morphological patterns of the intraparietal sulcus and the anterior intermediate parietal sulcus of Jensen in the human brain

Distinct parts of the intraparietal sulcal cortex contribute to sensorimotor integration and visual spatial attentional processing. A detailed examination of the morphological relations of the different segments of the complex intraparietal sulcal region in the human brain in standard stereotaxic space, which is a prerequisite for detailed structure-to-function studies, is not available. This study examined the intraparietal sulcus (IPS) and the related sulcus of Jensen in magnetic resonance imaging brain volumes registered in the Montreal Neurological Institute stereotaxic space. It was demonstrated that the IPS is divided into two branches: the anterior ramus and the posterior ramus of the IPS, often separated by a submerged gyral passage. The sulcus of Jensen emerges between the anterior and posterior rami of the IPS, and its ventral end is positioned between the first and second caudal branches of the superior temporal sulcus. In a small number of brains, the sulcus of Jensen may merge superficially with the first caudal branch of the superior temporal sulcus. The above morphological findings are discussed in relation to previously reported functional neuroimaging findings and provide the basis for future exploration of structure-to-function relations in the posterior parietal region of individual subjects.     

The generality of parietal involvement in visual attention.

Functional magnetic resonance imaging (fMRI) was used to determine whether different kinds of visual attention rely on a common neural substrate. Within one session, subjects performed three different attention experiments (each comparing an attentionally demanding task with an easier task using identical stimuli): (1) peripheral shifting, (2) object matching, and (3) a nonspatial conjunction task. Two areas were activated in all three experiments: one at the junction of intraparietal and transverse occipital sulci (IPTO), and another in the anterior intraparietal sulcus (AIPS). These regions are not simply involved in any effortful task, because they were not activated in a fourth experiment comparing a difficult language task with an easier control task. Thus, activity in IPTO and AIPS generalizes across a wide variety of attention-requiring tasks, supporting the existence of a common neural substrate underlying multiple modes of visual selection.     

Neural Activity during Natural Viewing of Sesame Street Statistically Predicts Test Scores in Early Childhood

It is not currently possible to measure the real-world thought process that a child has while observing an actual school lesson. However, if it could be done, children''s neural processes would presumably be predictive of what they know. Such neural measures would shed new light on children''s real-world thought. Toward that goal, this study examines neural processes that are evoked naturalistically, during educational television viewing. Children and adults all watched the same Sesame Street video during functional magnetic resonance imaging (fMRI). Whole-brain intersubject correlations between the neural timeseries from each child and a group of adults were used to derive maps of “neural maturity” for children. Neural maturity in the intraparietal sulcus (IPS), a region with a known role in basic numerical cognition, predicted children''s formal mathematics abilities. In contrast, neural maturity in Broca''s area correlated with children''s verbal abilities, consistent with prior language research. Our data show that children''s neural responses while watching complex real-world stimuli predict their cognitive abilities in a content-specific manner. This more ecologically natural paradigm, combined with the novel measure of “neural maturity,” provides a new method for studying real-world mathematics development in the brain.     

An fMRI Study of the Neural Systems Involved in Visually Cued Auditory Top-Down Spatial and Temporal Attention

Top-down attention to spatial and temporal cues has been thoroughly studied in the visual domain. However, because the neural systems that are important for auditory top-down temporal attention (i.e., attention based on time interval cues) remain undefined, the differences in brain activity between directed attention to auditory spatial location (compared with time intervals) are unclear. Using fMRI (magnetic resonance imaging), we measured the activations caused by cue-target paradigms by inducing the visual cueing of attention to an auditory target within a spatial or temporal domain. Imaging results showed that the dorsal frontoparietal network (dFPN), which consists of the bilateral intraparietal sulcus and the frontal eye field, responded to spatial orienting of attention, but activity was absent in the bilateral frontal eye field (FEF) during temporal orienting of attention. Furthermore, the fMRI results indicated that activity in the right ventrolateral prefrontal cortex (VLPFC) was significantly stronger during spatial orienting of attention than during temporal orienting of attention, while the DLPFC showed no significant differences between the two processes. We conclude that the bilateral dFPN and the right VLPFC contribute to auditory spatial orienting of attention. Furthermore, specific activations related to temporal cognition were confirmed within the superior occipital gyrus, tegmentum, motor area, thalamus and putamen.     

Cortical activations in humans grasp-related areas depend on hand used and handedness

Background

In non-human primates grasp-related sensorimotor transformations are accomplished in a circuit involving the anterior intraparietal sulcus (area AIP) and both the ventral and the dorsal sectors of the premotor cortex (vPMC and dPMC, respectively). Although a human homologue of such a circuit has been identified whether activity within this circuit varies depending on handedness has yet to be investigated.

Methodology/Principal Findings

We used functional magnetic resonance imaging (fMRI) to explicitly test how handedness modulates activity within human grasping-related brain areas. Right- and left-handers subjects were requested to reach towards and grasp an object with either the right or the left hand using a precision grip while scanned. A kinematic study was conducted with similar procedures as a behavioral counterpart for the fMRI experiment. Results from a factorial design revealed significant activity within the right dPMC, the right cerebellum and AIP bilaterally. The pattern of activity within these areas mirrored the results found for the behavioral study.

Conclusion/Significance

Data are discussed in terms of an handedness-independent role for the right dPMC in monitoring hand shaping, the need for bilateral AIP activity for the performance of precision grip movements which varies depending on handedness and the involvement of the cerebellum in terms of its connections with AIP. These results provide the first compelling evidence of specific grasping related neural activity depending on handedness.     

Form and motion coherence activate independent, but not dorsal/ventral segregated, networks in the human brain

There is much evidence in primates' visual processing for distinct mechanisms involved in object recognition and encoding object position and motion, which have been identified with 'ventral' and 'dorsal' streams, respectively, of the extra-striate visual areas [1] [2] [3]. This distinction may yield insights into normal human perception, its development and pathology. Motion coherence sensitivity has been taken as a test of global processing in the dorsal stream [4] [5]. We have proposed an analogous 'form coherence' measure of global processing in the ventral stream [6]. In a functional magnetic resonance imaging (fMRI) experiment, we found that the cortical regions activated by form coherence did not overlap with those activated by motion coherence in the same individuals. Areas differentially activated by form coherence included regions in the middle occipital gyrus, the ventral occipital surface, the intraparietal sulcus, and the temporal lobe. Motion coherence activated areas consistent with those previously identified as V5 and V3a, the ventral occipital surface, the intraparietal sulcus, and temporal structures. Neither form nor motion coherence activated area V1 differentially. Form and motion foci in occipital, parietal, and temporal areas were nearby but showed almost no overlap. These results support the idea that form and motion coherence test distinct functional brain systems, but that these do not necessarily correspond to a gross anatomical separation of dorsal and ventral processing streams.     

Tracking the mind's image in the brain II: transcranial magnetic stimulation reveals parietal asymmetry in visuospatial imagery

The functional relevance of brain activity during visuospatial tasks was investigated by combining functional magnetic resonance imaging with unilateral repetitive transcranial magnetic stimulation (rTMS). The cognitive tasks involved visuospatial operations on visually presented and mentally imagined material ("mental clock task"). While visuospatial operations were associated with activation of the intraparietal sulcus region bilaterally, only the group which received rTMS to the right parietal lobe showed an impairment of performance during and immediately after rTMS. This functional parietal asymmetry might indicate a capacity of the right parietal lobe to compensate for a temporary suppression of the left. This is compatible with current theories of spatial hemineglect and constitutes a constraint for models of distributed information processing in the parietal lobes.     

Spatiotemporal Dynamics of High-Gamma Activities during a 3-Stimulus Visual Oddball Task

Although many studies have investigated the neural basis of top-down and bottom-up attention, it still requires refinement in both temporal and spatial terms. We used magnetoencephalography to investigate the spatiotemporal dynamics of high-gamma (52–100 Hz) activities during top-down and bottom-up visual attentional processes, aiming to extend the findings from functional magnetic resonance imaging and event-related potential studies. Fourteen participants performed a 3-stimulus visual oddball task, in which both infrequent non-target and target stimuli were presented. We identified high-gamma event-related synchronization in the left middle frontal gyrus, the left intraparietal sulcus, the left thalamus, and the visual areas in different time windows for the target and non-target conditions. We also found elevated imaginary coherence between the left intraparietal sulcus and the right middle frontal gyrus in the high-gamma band from 300 to 400 ms in the target condition, and between the left thalamus and the left middle frontal gyrus in theta band from 150 to 450 ms. In addition, the strength of high-gamma imaginary coherence between the left middle frontal gyrus and left intraparietal sulcus, between the left middle frontal gyrus and the right middle frontal gyrus, and the high-gamma power in the left thalamus predicted inter-subject variation in target detection response time. This source-level electrophysiological evidence enriches our understanding of bi-directional attention processes: stimulus-driven bottom-up attention orientation to a salient, but irrelevant stimulus; and top-down allocation of attentional resources to stimulus evaluation.     

Neural Activity Changes Associated with Impulsive Responding in the Sustained Attention to Response Task

Humans can anticipate and prepare for uncertainties to achieve a goal. However, it is difficult to maintain this effort over a prolonged period of time. Inappropriate behavior is impulsively (or mindlessly) activated by an external trigger, which can result in serious consequences such as traffic crashes. Thus, we examined the neural mechanisms underlying such impulsive responding using functional magnetic resonance imaging (fMRI). Twenty-two participants performed a block-designed sustained attention to response task (SART), where each task block was composed of consecutive Go trials followed by a NoGo trial at the end. This task configuration enabled us to measure compromised preparation for NoGo trials during Go responses using reduced Go reaction times. Accordingly, parametric modulation analysis was conducted on fMRI data using block-based mean Go reaction times as an online marker of impulsive responding in the SART. We found that activity in the right dorsolateral prefrontal cortex (DLPFC) and the bilateral intraparietal sulcus (IPS) was positively modulated with mean Go reaction times. In addition, activity in the medial prefrontal cortex (MPFC) and the posterior cingulate cortex (PCC) was negatively modulated with mean Go reaction times, albeit statistically weakly. Taken together, spontaneously reduced activity in the right DLPFC and the IPS and spontaneously elevated activity in the MPFC and the PCC were associated with impulsive responding in the SART. These results suggest that such a spontaneous transition of brain activity pattern results in impulsive responding in monotonous situations, which in turn, might cause human errors in actual work environments.