Long-distance feedback projections to area V1: Implications for multisensory integration,spatial awareness,and visual consciousness

It is generally agreed that information flow through the cortex is constrained by a hierarchical architecture. Recent experimental evidence suggests that projections descending the hierarchy and targeting the primary visual cortex (area V1) may play an essential role in perceptual processes. We have, therefore, reexamined feedback projections to area V1, using retrograde tracer injections in this area. In addition to well-known areas, quantification of labeling in higher cortical areas reveals a number of hitherto unknown long-distance feedback connections originating from auditory (A1), multisensory (STP) cortices, but also from a perirhinal area (36). These feedback projections from advanced cortical stations, a global feature shared by areas that belong to the ventral visual stream, could play an important role in early multisensory integration and spatial awareness and could provide the physical substrate for the involvement of area V1 in visual consciousness.     

Atrophy of the parietal lobe in preclinical dementia

Cortical grey matter atrophy patterns have been reported in healthy ageing and Alzheimer disease (AD), but less consistently in the parietal regions of the brain. We investigated cortical grey matter volume patterns in parietal areas. The grey matter of the somatosensory cortex, superior and inferior parietal lobule was measured in 75 older adults (38 cognitively stable and 37 individuals with cognitive decline after 3 years). Dementia screening 6 years after scanning resulted in nine AD cases from the cognitively stable (n=3) and cognitive decline group (n=6), who were assigned to a third group, the preclinical AD group. When regional differences in cortical volume in the parietal lobe areas were compared between groups, significant differences were found between either the cognitive decline or stable group on the one hand and preclinical AD individuals on the other hand in the inferior parietal lobule. Group membership was best predicted by the grey matter volume of the inferior parietal lobule, compared to the other parietal lobe areas. The parietal lobe was characterised by a differential atrophy pattern based on cognitive status, which is in agreement with the 'last-developed-first-atrophied' principle. Future studies should investigate the surplus value of the inferior parietal lobe as a potential marker for the diagnosis of AD compared to other brain regions, such as the medial temporal lobe and the prefrontal lobe.     

Transfer of visual information after unilateral input to the brain

Visual callosal connections are more numerous and widespread in the association areas than in the primary visual cortex and adjoining visual areas. In keeping with this, the amount of "wrong" ipsilateral visual field that is represented in the various cortical areas of primates and cats increases as one goes from primary visual cortex to extraoccipital areas. Therefore it can be argued that transfer of unilaterally presented visual stimuli occurs mainly at the temporal and parietal cortical level.     

Large,colorful, or noisy? Attribute- and modality-specific activations during retrieval of perceptual attribute knowledge

Position emission tomography was used to investigate whether retrieval of perceptual knowledge from long-term memory activates unique cortical regions associated with the modality and/or attribute type retrieved. Knowledge about the typical color, size, and sound of common objects and animals was probed, in response to written words naming the objects. Relative to a nonsemantic control task, all the attribute judgments activated similar left temporal and frontal regions. Visual (color, size) knowledge selectively activated the right posterior inferior temporal (PIT) cortex, whereas sound judgments elicited selective activation in the left posterior superior temporal gyrus and the adjacent parietal cortex. All of the attribute judgments activated a left PIT region, but color retrieval generated more activation in this area. Size judgments activated the right medial parietal cortex. These results indicate that the retrieval of perceptual semantic information activates not only a general semantic network, but also cortical areas specialized for the modality and attribute type of the knowledge retrieved.     

Cortical localisation of the visual and auditory word form areas: a reconsideration of the evidence

In this paper we examine the evidence for human brain areas dedicated to visual or auditory word form processing by comparing cortical activation for auditory word repetition, reading, picture naming, and environmental sound naming. Both reading and auditory word repetition activated left lateralised regions in the frontal operculum (Broca's area), posterior superior temporal gyrus (Wernicke's area), posterior inferior temporal cortex, and a region in the mid superior temporal sulcus relative to baseline conditions that controlled for sensory input and motor output processing. In addition, auditory word repetition increased activation in a lateral region of the left mid superior temporal gyrus but critically, this area is not specific to auditory word processing, it is also activated in response to environmental sounds. There were no reading specific activations, even in the areas previously claimed as visual word form areas: activations were either common to reading and auditory word repetition or common to reading and picture naming. We conclude that there is no current evidence for cortical sites dedicated to visual or auditory word form processing.     

Timing of human cortical functions during cognition: role of MEG

Understanding of sensory and cognitive brain processes requires information about activation timing within and between different brain sites. Such data can be obtained by magnetoencephalography (MEG) that tracks cortical activation sequences with a millisecond temporal accuracy. MEG is gaining a well-established role in human neuroscience, complementing with its excellent temporal resolution the spatially more focused brain imaging methods. As examples of MEG's role in cognitive neuroscience, we discuss time windows related to cortical processing of sensory and multisensory stimuli, effects of the subject's own voice on the activity of their auditory cortex, timing of brain activation in reading, and cortical dynamics of the human mirror-neuron system activated when the subject views another person's movements.     

Neural correlates of rumination in adolescents with remitted major depressive disorder and healthy controls

The aim of the present study was to use fMRI to examine the neural correlates of engaging in rumination among a sample of remitted depressed adolescents, a population at high risk for future depressive relapse. A rumination induction task was used to assess differences in the patterns of neural activation during rumination versus a distraction condition among 26 adolescents in remission from major depressive disorder (rMDD) and in 15 healthy control adolescents. Self-report depression and rumination, as well as clinician-rated depression, were also assessed among all participants. All of the participants recruited regions in the default mode network (DMN), including the posterior cingulate cortex, medial prefrontal cortex, inferior parietal lobe, and medial temporal gyrus, during rumination. Increased activation in these regions during rumination was correlated with increased self-report rumination and symptoms of depression across all participants. Adolescents with rMDD also exhibited greater activation in regions involved in visual, somatosensory, and emotion processing than did healthy peers. The present findings suggest that during ruminative thought, adolescents with rMDD are characterized by increased recruitment of regions within the DMN and in areas involved in visual, somatosensory, and emotion processing.     

The specialized structure of human language cortex: pyramidal cell size asymmetries within auditory and language-associated regions of the temporal lobes

Functional lateralization of language within the cerebral cortex has long driven the search for structural asymmetries that might underlie language asymmetries. Most examinations of structural asymmetry have focused upon the gross size and shape of cortical regions in and around language areas. In the last 20 years several labs have begun to document microanatomical asymmetries in the structure of language-associated cortical regions. Such microanatomic results provide useful constraints and clues to our understanding of the biological bases of language specialization in the cortex. In a previous study we documented asymmetries in the size of a specific class of pyramidal cells in the superficial cortical layers. The present work uses a nonspecific stain for cell bodies to demonstrate the presence of an asymmetry in layer III pyramidal cell sizes within auditory, secondary auditory and language-associated regions of the temporal lobes. Specifically, the left hemisphere contains a greater number of the largest pyramidal cells, those that are thought to be the origin of long-range cortico-cortical connections. These results are discussed in the context of cortical columns and how such an asymmetry might alter cortical processing. These findings, in conjunction with other asymmetries in cortical organization that have been documented within several labs, clearly demonstrate that the columnar and connective structure of auditory and language cortex in the left hemisphere is distinct from homotopic regions in the contralateral hemisphere.     

The cortical language circuit: from auditory perception to sentence comprehension

Over the years, a large body of work on the brain basis of language comprehension has accumulated, paving the way for the formulation of a comprehensive model. The model proposed here describes the functional neuroanatomy of the different processing steps from auditory perception to comprehension as located in different gray matter brain regions. It also specifies the information flow between these regions, taking into account white matter fiber tract connections. Bottom-up, input-driven processes proceeding from the auditory cortex to the anterior superior temporal cortex and from there to the prefrontal cortex, as well as top-down, controlled and predictive processes from the prefrontal cortex back to the temporal cortex are proposed to constitute the cortical language circuit.     

Cerebellar output: motor and cognitive channels

The input to the cerebellum has long been known to originate from widespread regions of the cerebral cortex including the frontal, parietal and temporal lobes. The output of the cerebellum, however, was thought to project mainly to the primary motor cortex. Recent anatomical observations have challenged this view. It is now apparent that cerebellar output goes to multiple cortical areas, including not only the primary motor cortex, but also areas of premotor and prefrontal cortex. In fact, there is growing evidence that each of the areas of cerebral cortex that project to the cerebellum is also the target of cerebellar output. The cerebellar output to individual cortical areas originates from distinct clusters of neurons in the deep nuclei which we have termed `output channels'. The individual output channels to the cortical areas we have examined display little or no overlap. Physiological recordings in awake trained primates indicate that neurons in different output channels appear to be involved in distinct aspects of behavior, and in both motor and cognitive functions. These observations indicate that the cerebellar influence on the cerebral cortex is more extensive than previously recognized.     

Multisensory integration in multiple reference frames in the posterior parietal cortex

Spatial information processing takes place in different brain regions that receive converging inputs from several sensory modalities. Because of our own movements—for example, changes in eye position, head rotations, and so forth—unimodal sensory representations move continuously relative to one another. It is generally assumed that for multisensory integration to be an orderly process, it should take place between stimuli at congruent spatial locations. In the monkey posterior parietal cortex, the ventral intraparietal (VIP) area is specialized for the analysis of movement information using visual, somatosensory, vestibular, and auditory signals. Focusing on the visual and tactile modalities, we found that in area VIP, like in the superior colliculus, multisensory signals interact at the single neuron level, suggesting that this area participates in multisensory integration. Curiously, VIP does not use a single, invariant coordinate system to encode locations within and across sensory modalities. Visual stimuli can be encoded with respect to the eye, the head, or halfway between the two reference frames, whereas tactile stimuli seem to be prevalently encoded relative to the body. Hence, while some multisensory neurons in VIP could encode spatially congruent tactile and visual stimuli independently of current posture, in other neurons this would not be the case. Future work will need to evaluate the implications of these observations for theories of optimal multisensory integration.Edited by: Marie-Hélène Giard and Mark Wallace     

Information-processing modules and their relative modality specificity

This research uses fMRI to understand the role of eight cortical regions in a relatively complex information-processing task. Modality of input (visual versus auditory) and modality of output (manual versus vocal) are manipulated. Two perceptual regions (auditory cortex and fusiform gyrus) only reflected perceptual encoding. Two motor regions were involved in information rehearsal as well as programming of overt actions. Two cortical regions (parietal and prefrontal) performed processing (retrieval and representational change) independent of input and output modality. The final two regions (anterior cingulate and caudate) were involved in control of cognition independent of modality of input or output and content of the material. An information-processing model, based on the ACT-R theory, is described that predicts the BOLD response in these regions. Different modules in the theory vary in the degree to which they are modality-specific and the degree to which they are involved in central versus peripheral cognitive processes.     

Prefrontal cortex regulates inhibition and excitation in distributed neural networks.

Prefrontal cortex provides both inhibitory and excitatory input to distributed neural circuits required to support performance in diverse tasks. Neurological patients with prefrontal damage are impaired in their ability to inhibit task-irrelevant information during behavioral tasks requiring performance over a delay. The observed enhancements of primary auditory and somatosensory cortical responses to task-irrelevant distractors suggest that prefrontal damage disrupts inhibitory modulation of inputs to primary sensory cortex, perhaps through abnormalities in a prefrontal-thalamic sensory gating system. Failure to suppress irrelevant sensory information results in increased neural noise, contributing to the deficits in decision making routinely observed in these patients. In addition to a critical role in inhibitory control of sensory flow to primary cortical regions, and tertiary prefrontal cortex also exerts excitatory input to activity in multiple sub-regions of secondary association cortex. Unilateral prefrontal damage results in multi-modal decreases in neural activity in posterior association cortex in the hemisphere ipsilateral to damage. This excitatory modulation is necessary to sustain neural activity during working memory. Thus, prefrontal cortex is able to sculpt behavior through parallel inhibitory and excitatory regulation of neural activity in distributed neural networks.     

The supramodal brain: implications for auditory perception

ABSTRACT

The perceptual brain is designed around multisensory input. Areas once thought dedicated to a single sense are now known to work with multiple senses. It has been argued that the multisensory nature of the brain reflects a cortical architecture for which task, rather than sensory system, is the primary design principle. This supramodal thesis is supported by recent research on human echolocation and multisensory speech perception. In this review, we discuss the behavioural implications of a supramodal architecture, especially as they pertain to auditory perception. We suggest that the architecture implies a degree of perceptual parity between the senses and that cross-sensory integration occurs early and completely. We also argue that a supramodal architecture implies that perceptual experience can be shared across modalities and that this sharing should occur even without bimodal experience. We finish by briefly suggesting areas of future research.     

Listening to polyphonic music recruits domain-general attention and working memory circuits

Polyphonic music combines multiple auditory streams to create complex auditory scenes, thus providing a tool for investigating the neural mechanisms that orient attention in natural auditory contexts. Across two fMRI experiments, we varied stimuli and task demands in order to identify the cortical areas that are activated during attentive listening to real music. In individual experiments and in a conjunction analysis of the two experiments, we found bilateral blood oxygen level dependent (BOLD) signal increases in temporal (the superior temporal gyrus), parietal (the intraparietal sulcus), and frontal (the precentral sulcus, the inferior frontal sulcus and gyrus, and the frontal operculum) areas during selective and global listening, as compared with passive rest without musical stimulation. Direct comparisons of the listening conditions showed significant differences between attending to single timbres (instruments) and attending across multiple instruments, although the patterns that were observed depended on the relative demands of the tasks being compared. The overall pattern of BOLD signal increases indicated that attentive listening to music recruits neural circuits underlying multiple forms of working memory, attention, semantic processing, target detection, and motor imagery. Thus, attentive listening to music appears to be enabled by areas that serve general functions, rather than by music-specific cortical modules.     

Investigating the modality specificity of response selection using a temporal flanker task

The neurocognitive architecture for response selection is uncertain. Some theorists suggest that it is mediated by an amodal central mechanism, whereas others propose a set of independent control mechanisms. In a functional neuroimaging experiment, we investigated the nature of response selection by examining how its underlying brain mechanisms are affected by stimulus modality. To do this, we used a modified flanker task, in which the target and flanker (distractor) stimuli differed in time rather than space, making it accessible for both visual and auditory stimuli. As in the traditional flanker task, larger reaction times were observed for incongruent than congruent trials (i.e., a congruency effect) for both modalities. Congruency affected brain activation for both modalities in prefrontal cortex, parietal cortex, and the putamen. Modality-dependent activation was found in additional prefrontal and parietal regions for the visual modality and in left inferior prefrontal cortex for the auditory modality. Modality-dependent activity specifically related to response congruency was also found in sensory cortical regions. These data suggest that modality affects the brain regions throughout the cortex mediating response selection even for conceptually identical stimuli and tasks. They are consistent with the hypothesis that (at least partially) independent brain networks mediate response selection and that input modality may be a powerful factor for organizing neural activity to support task performance.     

Cortical dynamics of contextually cued attentive visual learning and search: spatial and object evidence accumulation

How do humans use target-predictive contextual information to facilitate visual search? How are consistently paired scenic objects and positions learned and used to more efficiently guide search in familiar scenes? For example, humans can learn that a certain combination of objects may define a context for a kitchen and trigger a more efficient search for a typical object, such as a sink, in that context. The ARTSCENE Search model is developed to illustrate the neural mechanisms of such memory-based context learning and guidance and to explain challenging behavioral data on positive-negative, spatial-object, and local-distant cueing effects during visual search, as well as related neuroanatomical, neurophysiological, and neuroimaging data. The model proposes how global scene layout at a first glance rapidly forms a hypothesis about the target location. This hypothesis is then incrementally refined as a scene is scanned with saccadic eye movements. The model simulates the interactive dynamics of object and spatial contextual cueing and attention in the cortical What and Where streams starting from early visual areas through medial temporal lobe to prefrontal cortex. After learning, model dorsolateral prefrontal cortex (area 46) primes possible target locations in posterior parietal cortex based on goal-modulated percepts of spatial scene gist that are represented in parahippocampal cortex. Model ventral prefrontal cortex (area 47/12) primes possible target identities in inferior temporal cortex based on the history of viewed objects represented in perirhinal cortex.     

Multisensory cortical processing of object shape and its relation to mental imagery

Here, we used functional magnetic resonance imaging to investigate the multisensory processing of object shape in the human cerebral cortex and explored the role of mental imagery in such processing. Regions active bilaterally during both visual and haptic shape perception, relative to texture perception in the respective modality, included parts of the superior parietal gyrus, the anterior intraparietal sulcus, and the lateral occipital complex. Of these bimodal regions, the lateral occipital complexes preferred visual over haptic stimuli, whereas the parietal areas preferred haptic over visual stimuli. Whereas most subjects reported little haptic imagery during visual shape perception, experiences of visual imagery during haptic shape perception were common. Across subjects, ratings of the vividness of visual imagery strongly predicted the amount of haptic shape-selective activity in the right, but not in the left, lateral occipital complex. Thus, visual imagery appears to contribute to activation of some, but not all, visual cortical areas during haptic perception.     

Episodic retrieval and the cortical binding of relational activity

Retrieval of episodic memories depends on the successful “re-collection” of event features, such as the time, place, people, thoughts, and feelings associated with a past experience.In neuroimaging studies, ventral regions of the posterior parietal cortex (vPPC) are particularly active when episodic memories are successfully retrieved. A review of the neural correlates of episodic retrieval is presented along with a new theory, cortical binding of relational activity (CoBRA). According to CoBRA, the vPPC acts as a convergence zone that binds episodic features stored in disparate neocortical regions. This process works in conjunction with other known mechanisms, such as those associated with the prefrontal cortex and medial temporal lobe.