Neural mechanisms underlying reaching for remembered targets cued kinesthetically or visually in left or right hemispace

Reaching for a target involves integrative coordinate transformation processes between the representation of the target location, the sensorimotor information of limb of reach, and body space. Although right hemisphere dominance for visuospatial information processing is well established, corresponding right hemisphere dominance for kinesthetic spatial information processing remains to be demonstrated. We explored neural mechanisms of encoding target locations using 15O-butanol positron emission tomography (PET) in normal volunteers in a factorial experiment, where modality (visual/kinesthetic) and hemispace of target presentation (left/right of midsagittal plane) were varied systematically. After target presentation, subjects reached to the encoded target location. PET data analysis using SPM99 showed increased neural activity (P < 0.05, corrected) associated with left hemispace target presentation in right hemisphere areas (sensorimotor, anterior cingulate, insular, and temporo-occipital cortex) only. By contrast, right hemispace target presentation activated bilateral temporo-occipital cortex, which extended into the right temporo-parietal cortex and left sensorimotor cortex. A significant interaction of hemispace and modality of target presentation observed in right temporo-parietal cortex resulted from an increase in neural activity with kinesthetic target presentation in right hemispace. The data support an important role for the right temporo-parietal area in visuospatial processing and suggest a specific role of the right hemisphere in kinesthetic spatial processing.     

Executive control of spatial attention shifts in the auditory compared to the visual modality

Voluntary orienting of visual spatial attention has been shown to be associated with activation in a distributed network of frontal and parietal brain areas. Neuropsychological data suggest that at least some of these areas should be sensitive to the direction in which attention is shifted. The aim of this study was to use rapid event-related functional magnetic resonance imaging to investigate whether spatial attention in the auditory modality is subserved by the same or different brain areas as in the visual modality, and whether the auditory and visual attention networks show any degree of hemispheric lateralisation or sensitivity to the direction of attention shifts. The results suggest that auditory and visual spatial attention shifts are controlled by a supramodal network of frontal, parietal and temporal areas. Areas activated included the precuneus and superior parietal cortex, the inferior parietal cortex and temporo-parietal junction, as well as the premotor and supplementary motor areas and dorsolateral prefrontal cortex (DLPFC). In the auditory task, some of these areas, in particular the precuneus as well as the inferior parietal cortex and temporo-parietal junction, showed 'relative' asymmetry, in that they responded more strongly to attention shifts towards the contralateral than the ipsilateral hemispace. Some areas, such as the right superior parietal cortex and left DLPFC, showed 'absolute' asymmetry, in that they responded more strongly in one than in the other hemisphere.     

The cortical substrate of visual extinction

Neuroimaging studies investigated the attentional systems of the human brain revealing two networks, one for voluntary allocation of attention and another for stimulus-driven attentional processes. Whereas lesions of the latter system were supposed to lead to spatial neglect, we show that such lesions rather are typical for the occurrence of visual extinction. Extinction describes the inability of brain-damaged patients to detect a contralesional target in the presence of a competing ipsilesional stimulus. In a sample of consecutively admitted patients with right hemisphere stroke, we found dissociable cortical substrates for spatial neglect and visual extinction. There was a surprising congruency between the typical lesion site in patients with extinction and the activation clusters found in previous neuroimaging studies of healthy subjects. The results show that the temporo-parietal junction (TPJ), considered to be a crucial part of the stimulus-driven attentional network, is the neural substrate of visual extinction.     

Neural systems for visual orienting and their relationships to spatial working memory

We investigated neural correlates of human visual orienting using event-related functional magnetic resonance imaging (fMRI). When subjects voluntarily directed attention to a peripheral location, we recorded robust and sustained signals uniquely from the intraparietal sulcus (IPs) and superior frontal cortex (near the frontal eye field, FEF). In the ventral IPs and FEF only, the blood oxygen level dependent signal was modulated by the direction of attention. The IPs and FEF also maintained the most sustained level of activation during a 7-sec delay, when subjects maintained attention at the peripheral cued location (working memory). Therefore, the IPs and FEF form a dorsal network that controls the endogenous allocation and maintenance of visuospatial attention. A separate right hemisphere network was activated by the detection of targets at unattended locations. Activation was largely independent of the target's location (visual field). This network included among other regions the right temporo-parietal junction and the inferior frontal gyrus. We propose that this cortical network is important for reorienting to sensory events.     

Identification of language-specific brain activity using magnetoencephalography

The purpose of the present investigation was to explore the ability of magnetoencephalography (MEG) to identify brain areas involved in language comprehension. Event-related magnetic fields (ERFs) were recorded from 7 right-handed adults with no history of neurological disorder or learning disability as they engaged in an auditory and a visual word-recognition task. A face-recognition task served as control. During the later portion of the ERFs, activity sources from both language tasks tended to overlap in temporal and temporo-parietal cortices. There was a clear preponderance of such sources in the left compared to the right hemisphere in all participants. These findings demonstrate that MEG is a promising tool for identifying brain regions involved in the analysis of linguistic stimuli, in addition to the initial encoding of stimulus features.     

Identification of Language-Specific Brain Activity Using Magnetoencephalography

The purpose of the present investigation was to explore the ability of magnetoencephalography (MEG) to identify brain areas involved in language comprehension. Event-related magnetic fields (ERFs) were recorded from 7 right-handed adults with no history of neurological disorder or learning disability as they engaged in an auditory and a visual word-recognition task. A face-recognition task served as control. During the later portion of the ERFs, activity sources from both language tasks tended to overlap in temporal and temporo-parietal cortices. There was a clear preponderance of such sources in the left compared to the right hemisphere in all participants. These findings demonstrate that MEG is a promising tool for identifying brain regions involved in the analysis of linguistic stimuli, in addition to the initial encoding of stimulus features.     

Attentional bias to briefly presented emotional distractors follows a slow time course in visual cortex

A central controversy in the field of attention is how the brain deals with emotional distractors and to what extent they capture attentional processing resources reflexively due to their inherent significance for guidance of adaptive behavior and survival. Especially, the time course of competitive interactions in early visual areas and whether masking of briefly presented emotional stimuli can inhibit biasing of processing resources in these areas is currently unknown. We recorded frequency-tagged potentials evoked by a flickering target detection task in the foreground of briefly presented emotional or neutral pictures that were followed by a mask in human subjects. We observed greater competition for processing resources in early visual cortical areas with shortly presented emotional relative to neutral pictures ~275 ms after picture offset. This was paralleled by a reduction of target detection rates in trials with emotional pictures ~400 ms after picture offset. Our finding that briefly presented emotional distractors are able to bias attention well after their offset provides evidence for a rather slow feedback or reentrant neural competition mechanism for emotional distractors that continues after the offset of the emotional stimulus.     

Neural correlates of conscious and unconscious vision in parietal extinction

Brain areas activated by stimuli in the left visual field of a right parietal patient suffering from left visual extinction were identified using event-related functional magnetic resonance imaging. Left visual field stimuli that were extinguished from awareness still activated the ventral visual cortex, including areas in the damaged right hemisphere. An extinguished face stimulus on the left produced robust category-specific activation of the right fusiform face area. On trials where the left visual stimulus was consciously seen rather than extinguished, greater activity was found in the ventral visual cortex of the damaged hemisphere, and also in frontal and parietal areas of the intact hemisphere. These findings extend recent observations on visual extinction, suggesting distinct neural correlates for conscious and unconscious perception.     

Neural Correlates of Conscious and Unconscious Vision in Parietal Extinction

Brain areas activated by stimuli in the left visual field of a right parietal patient suffering from left visual extinction were identified using event-related functional magnetic resonance imaging. Left visual field stimuli that were extinguished from awareness still activated the ventral visual cortex, including areas in the damaged right hemisphere. An extinguished face stimulus on the left produced robust category-specific activation of the right fusiform face area. On trials where the left visual stimulus was consciously seen rather than extinguished, greater activity was found in the ventral visual cortex of the damaged hemisphere, and also in frontal and parietal areas of the intact hemisphere. These findings extend recent observations on visual extinction, suggesting distinct neural correlates for conscious and unconscious perception.     

Parietal blood oxygenation level‐dependent response evoked by covert visual search reflects set‐size effect in monkeys

Distinguishing a target from distractors during visual search is crucial for goal‐directed behaviour. The more distractors that are presented with the target, the larger is the subject's error rate. This observation defines the set‐size effect in visual search. Neurons in areas related to attention and eye movements, like the lateral intraparietal area (LIP) and frontal eye field (FEF), diminish their firing rates when the number of distractors increases, in line with the behavioural set‐size effect. Furthermore, human imaging studies that have tried to delineate cortical areas modulating their blood oxygenation level‐dependent (BOLD) response with set size have yielded contradictory results. In order to test whether BOLD imaging of the rhesus monkey cortex yields results consistent with the electrophysiological findings and, moreover, to clarify if additional other cortical regions beyond the two hitherto implicated are involved in this process, we studied monkeys while performing a covert visual search task. When varying the number of distractors in the search task, we observed a monotonic increase in error rates when search time was kept constant as was expected if monkeys resorted to a serial search strategy. Visual search consistently evoked robust BOLD activity in the monkey FEF and a region in the intraparietal sulcus in its lateral and middle part, probably involving area LIP. Whereas the BOLD response in the FEF did not depend on set size, the LIP signal increased in parallel with set size. These results demonstrate the virtue of BOLD imaging in monkeys when trying to delineate cortical areas underlying a cognitive process like visual search. However, they also demonstrate the caution needed when inferring neural activity from BOLD activity.     

A neural network elicited by parametric manipulation of the attention load

We used a parametric experimental design to identify the rCBF variations related to a continuous variation of the attention load. The experiment involved goal-directed visual tasks. The length of time during which the subject's attention was engaged toward the external stimulus was taken as the factor of interest. The neural network revealed areas that positively (left cerebellum, bilateral MT/V5 complex and superior parietal lobule, right inferior temporal lobe and dorsolateral prefrontal cortex) or negatively (precuneus, anterior cingulate and medial superior frontal cortex) correlate with the attention load. Results demonstrate that the activity of these areas varies continuously as a function of the variation in the attention load.     

Processing of auditory spatial cues in human cortex: an fMRI study

The issue of where in the human cortex coding of sound location is represented still is a matter of debate. It is unclear whether there are cortical areas that are specifically activated depending on the location of sound. Are identical or distinct cortical areas in one hemisphere involved in processing of sounds from the left and right? Also, the possibility has not been investigated so far that distinct areas have a preference for processing of central and eccentric sound locations. The present study focussed on these issues by using functional magnetic resonance imaging (fMRI). Activations evoked by left, right and central sounds were analysed separately, and contrasts were computed between these conditions. We did not find areas, which were involved in the processing of exclusively left, right or central sound positions. Large overlapping areas rather were observed for the three sound stimuli, located in the temporal, parietal and frontal cortices of both hemispheres. This result argues for the idea of a widely distributed bilateral network accessing an internal representation of the body to encode stimulus position in relation to the body median plane. However, two areas (right BA 40 and left BA 37) also were found to have preferences for sound position. In particular, BA 40 turned out to be significantly more activated by processing central positions, compared to eccentric stimuli. In line with previous findings on visual perception, the latter observation supports the assumption that the right inferior parietal cortex may be preferentially involved in the perception of central stimulus positions in relation to the body.     

Neural blackboard architectures: the realization of compositionality and systematicity in neural networks

In this paper, we will first introduce the notions of systematicity and combinatorial productivity and we will argue that these notions are essential for human cognition and probably for every agent that needs to be able to deal with novel, unexpected situations in a complex environment. Agents that use compositional representations are faced with the so-called binding problem and the question of how to create neural network architectures that can deal with it is essential for understanding higher level cognition. Moreover, an architecture that can solve this problem is likely to scale better with problem size than other neural network architectures. Then, we will discuss object-based attention. The influence of spatial attention is well known, but there is solid evidence for object-based attention as well. We will discuss experiments that demonstrate object-based attention and will discuss a model that can explain the data of these experiments very well. The model strongly suggests that this mode of attention provides a neural basis for parallel search. Next, we will show a model for binding in visual cortex. This model is based on a so-called neural blackboard architecture, where higher cortical areas act as processors, specialized for specific features of a visual stimulus, and lower visual areas act as a blackboard for communication between these processors. This implies that lower visual areas are involved in more than bottom-up visual processing, something which already was apparent from the large number of recurrent connections from higher to lower visual areas. This model identifies a specific role for these feedback connections. Finally, we will discuss the experimental evidence that exists for this architecture.     

Brain regions underlying repetition and auditory-verbal short-term memory deficits in aphasia: Evidence from voxel-based lesion symptom mapping

Background: A deficit in the ability to repeat auditory-verbal information is common among individuals with aphasia. The neural basis of this deficit has traditionally been attributed to the disconnection of left posterior and anterior language regions via damage to a white matter pathway, the arcuate fasciculus. However, a number of lesion and imaging studies have called this notion into question.

Aims: The goal of this study was to identify the neural correlates of repetition and a related process, auditory-verbal short-term memory (AVSTM). Both repetition and AVSTM involve common elements such as auditory and phonological analysis and translation to speech output processes. Based on previous studies, we predicted that both repetition and AVSTM would be most dependent on posterior language regions in left temporo-parietal cortex.

Methods & Procedures: We tested 84 individuals with left hemisphere lesions due to stroke on an experimental battery of repetition and AVSTM tasks. Participants were tested on word, pseudoword, and number-word repetition, as well as digit and word span tasks. Brain correlates of these processes were identified using a statistical, lesion analysis approach known as voxel-based lesion symptom mapping (VLSM). VLSM allows for a voxel-by-voxel analysis of brain areas most critical to performance on a given task, including both grey and white matter regions.

Outcomes & Results: The VLSM analyses showed that left posterior temporo-parietal cortex, not the arcuate fasciculus, was most critical for repetition as well as for AVSTM. The location of maximal foci, defined as the voxels with the highest t values, varied somewhat among measures: Word and pseudoword repetition had maximal foci in the left posterior superior temporal gyrus, on the border with inferior parietal cortex, while word and digit span, as well as number-word repetition, were centred on the border between the middle temporal and superior temporal gyri and the underlying white matter.

Conclusions: Findings from the current study show that (1) repetition is most critically mediated by cortical regions in left posterior temporo-parietal cortex; (2) repetition and AVSTM are mediated by partially overlapping networks; and (3) repetition and AVSTM deficits can be observed in different types of aphasia, depending on the site and extent of the brain injury. These data have implications for the prognosis of chronic repetition and AVSTM deficits in individuals with aphasia when lesions involve critical regions in left temporo-parietal cortex.     

Neural correlates of self-focused attention in social anxiety

Socially anxious individuals tend to shift their attention away from external socially threatening cues and instead become highly self-focused. Such heightened self-focused attention has been suggested to be involved in the development and maintenance of social anxiety disorder. This study used functional magnetic resonance imaging to investigate the neural correlates of self-focused attention in 16 high socially anxious (HSA) and 16 low socially anxious (LSA) individuals. Participants were instructed to focus their attention either inwardly or outwardly during a simulated social situation. Results indicate hyperactivation of medial prefrontal cortex (mPFC), temporo-parietal junction (TPJ) and temporal pole during inward vs outward attention in HSA compared with LSA participants. Furthermore, activation of mPFC, right anterior insula, TPJ and posterior cingulate cortex was positively correlated with the trait of self-focused attention in HSA subjects. Results highlight the prominent role of the mPFC and other cortical structures in abnormal self-focused attention in social anxiety. Finally, findings for the insula suggest increased processing of bodily states that is related to the amount of habitual self-focused attention in social anxiety.     

Comparing the effects of auditory deprivation and sign language within the auditory and visual cortex

To investigate neural plasticity resulting from early auditory deprivation and use of American Sign Language, we measured responses to visual stimuli in deaf signers, hearing signers, and hearing nonsigners using functional magnetic resonance imaging. We examined "compensatory hypertrophy" (changes in the responsivity/size of visual cortical areas) and "cross-modal plasticity" (changes in auditory cortex responses to visual stimuli). We measured the volume of early visual areas (V1, V2, V3, V4, and MT+). We also measured the amplitude of responses within these areas, and within the auditory cortex, to a peripheral visual motion stimulus that was attended or ignored. We found no major differences between deaf and hearing subjects in the size or responsivity of early visual areas. In contrast, within the auditory cortex, motion stimuli evoked significant responses in deaf subjects, but not in hearing subjects, in a region of the right auditory cortex corresponding to Brodmann's areas 41, 42, and 22. This hemispheric selectivity may be due to a predisposition for the right auditory cortex to process motion; earlier studies report a right hemisphere bias for auditory motion in hearing subjects. Visual responses within the auditory cortex of deaf subjects were stronger for attended than ignored stimuli, suggesting top-down processes. Hearing signers did not show visual responses in the auditory cortex, indicating that cross-modal plasticity can be attributed to auditory deprivation rather than sign language experience. The largest effects of auditory deprivation occurred within the auditory cortex rather than the visual cortex, suggesting that the absence of normal input is necessary for large-scale cortical reorganization to occur.     

Hemispheric lateralization in top‐down attention during spatial relation processing: a Granger causal model approach

Magnetoencephalography was recorded during a matching‐to‐sample plus cueing paradigm, in which participants judged the occurrence of changes in either categorical (CAT) or coordinate (COO) spatial relations. Previously, parietal and frontal lobes were identified as key areas in processing spatial relations and it was shown that each hemisphere was differently involved and modulated by the scope of the attention window (e.g. a large and small cue). In this study, Granger analysis highlighted the patterns of causality among involved brain areas – the direction of information transfer ran from the frontal to the visual cortex in the right hemisphere, whereas it ran in the opposite direction in the left side. Thus, the right frontal area seems to exert top‐down influence, supporting the idea that, in this task, top‐down signals are selectively related to the right side. Additionally, for CAT change preceded by a small cue, the right frontal gyrus was not involved in the information transfer, indicating a selective specialization of the left hemisphere for this condition. The present findings strengthen the conclusion of the presence of a remarkable hemispheric specialization for spatial relation processing and illustrate the complex interactions between the lateralized parts of the neural network. Moreover, they illustrate how focusing attention over large or small regions of the visual field engages these lateralized networks differently, particularly in the frontal regions of each hemisphere, consistent with the theory that spatial relation judgements require a fronto‐parietal network in the left hemisphere for categorical relations and on the right hemisphere for coordinate spatial processing.     

Visuospatial processing in children with neurofibromatosis type 1

Neuroimaging studies investigating the neural network of visuospatial processing have revealed a right hemisphere network of activation including inferior parietal lobe, dorsolateral prefrontal cortex, and extrastriate regions. Impaired visuospatial processing, indicated by the Judgment of Line Orientation (JLO), is commonly seen in individuals with neurofibromatosis type 1 (NF-1). Nevertheless, few studies have examined the neural activity associated with visuospatial processing in NF-1, in particular, during a JLO task. This study used functional neuroimaging to explore differences in volume of activation in predefined regions of interest between 13 individuals with NF-1 and 13 controls while performing an analogue JLO task. We hypothesized that participants with NF-1 would show anomalous right hemisphere activation and therefore would recruit regions within the left hemisphere to complete the task. Multivariate analyses of variance were used to test for differences between groups in frontal, temporal, parietal, and occipital regions. Results indicate that, as predicted, controls utilized various right hemisphere regions to complete the task, while the NF-1 group tended to recruit left hemisphere regions. These results suggest that the NF-1 group has an inefficient right hemisphere network. An additional unexpected finding was that the NF-1 group showed decreased volume of activation in primary visual cortex (BA 17). Future studies are needed to examine whether the decrease in primary visual cortex is related to a deficit in basic visual processing; findings could ultimately lead to a greater understanding of the nature of deficits in NF-1 and have implications for remediation.     

Neural mechanisms of memory retrieval: role of the prefrontal cortex

In the primate brain, long-term memory is stored in the neocortical association area which is also engaged in sensory perception. The coded representation of memory is retrieved via interactions of hierarchically different cortical areas along bottom-up and top-down anatomical connections. The functional significance of the fronto-cortical top-down neuronal projections has been relevantly assessed in a new experimental paradigm using posterior-split-brain monkeys. When the splenium of the corpus callosum and the anterior commissure were selectively split, the bottom-up visual signal originating from the unilateral striate cortex could not reach the contralateral visual cortical areas. In this preparation, long-term memory acquired through visual stimulus-stimulus association learning was prevented from transferring across hemispheres. Nonetheless, following the presentation of a visual cue to one hemisphere, the prefrontal cortex could instruct the contralateral hemisphere to retrieve the correct stimulus specified by the cue. These results support the hypothesis that the prefrontal cortex can regulate memory recall in the absence of bottom-up sensory input. In humans, functional neuroimaging studies have revealed activation of a distributed neural network, including the prefrontal cortex, during memory retrieval tasks. Thus, the prefrontal cortex is consistently involved in retrieval of long-term memory in primates.     

Dissociation of attention and intention in human posterior parietal cortex: an fMRI study

In natural environments, the orienting of attention to an object of interest occurs jointly with selecting it as a potential target for action. This coupling of perceptual selection and motor planning has led to 'the premotor theory of attention', which argues that attention and intention share the same neural mechanism. Here we used fMRI to test this hypothesis by examining neural activity in the temporal parietal junction (TPJ) and intraparietal sulcus (IPS) of the posterior parietal cortex (PPC) while subjects searched for a target among distractors with the presence of color singletons. Task-relevant salience of stimuli and the variability of response time were used to characterize the behavior of attention and intention components of the visuomotor transformation performed in these two cortical regions, respectively. We found that TPJ responses were significantly higher when the color singletons were distractors (vs. targets), suggesting that the TPJ was involved in attentional control of interference from task-irrelevant but salient stimuli. In contrast, signals in the IPS were closely related to the variability of response time, with a larger BOLD response associated with longer RTs, suggesting that the IPS plays a pivotal role in intention by translating encoded information into action through evidence accumulation. Our data help to specify the functional division of labor between the IPS and TPJ and to further dissociate process components in visual search.