Neural substrates of tactile object recognition: an fMRI study

A functional magnetic resonance imaging (fMRI) study was conducted during which seven subjects carried out naturalistic tactile object recognition (TOR) of real objects. Activation maps, conjunctions across subjects, were compared between tasks involving TOR of common real objects, palpation of "nonsense" objects, and rest. The tactile tasks involved similar motor and sensory stimulation, allowing higher tactile recognition processes to be isolated. Compared to nonsense object palpation, the most prominent activation evoked by TOR was in secondary somatosensory areas in the parietal operculum (SII) and insula, confirming a modality-specific path for TOR. Prominent activation was also present in medial and lateral secondary motor cortices, but not in primary motor areas, supporting the high level of sensory and motor integration characteristic of object recognition in the tactile modality. Activation in a lateral occipitotemporal area associated previously with visual object recognition may support cross-modal collateral activation. Finally, activation in medial temporal and prefrontal areas may reflect a common final pathway of modality-independent object recognition. This study suggests that TOR involves a complex network including parietal and insular somatosensory association cortices, as well as occipitotemporal visual areas, prefrontal, and medial temporal supramodal areas, and medial and lateral secondary motor cortices. It confirms the involvement of somatosensory association areas in the recognition component of TOR, and the existence of a ventrolateral somatosensory pathway for TOR in intact subjects. It challenges the results of previous studies that emphasize the role of visual cortex rather than somatosensory association cortices in higher-level somatosensory cognition.     

The neural basis for simulated drawing and the semantic implications

This functional magnetic resonance imaging (fMRI) study of the mental simulation of drawing (1) investigated the neural substrates of drawing and (2) delineated the semantic aspects of drawing. The goal was to advance our understanding of how drawing a familiar object is linked to lexical semantics and therefore a viable method to use to rehabilitate aphasia. We hypothesized that the semantic aspects of drawing familiar objects compared to drawing non-objects would yield greater activation in the inferior temporal cortex and the inferior frontal cortex of the left hemisphere. To test this hypothesis, eight right-handed subjects performed an fMRI experiment that directly contrasted drawing familiar objects to non-objects using mental imagery. Simulated drawing recruited a large, distributed network of frontal, parietal, and temporal structures. In the contrast comparing drawing familiar objects to non-objects there was stronger activation in the left hemisphere within the inferior temporal, anterior inferior frontal, inferior parietal and superior frontal cortices. The activation within the inferior temporal cortex was associated with visual semantic processing and semantic mediated naming. We suggest that the anterior inferior frontal activation is linked to the inferior temporal cortex and is involved in the selection of specific semantic features of the object as well as retrieval of information regarding the perceptual aspects of the object.     

Neural substrates of tactile imagery: a functional MRI study

fMRI investigation on the neural substrates involved in tactile imagery is reported. Healthy subjects performed mental imagery of tactile stimulation on the dorsal aspect of the right hand. The results were compared with the regions of activation during the actual tactile stimulation. During imagery, contralateral primary and secondary somatosensory areas were activated along with activation in the left parietal lobe. Activations in left inferior frontal gyri (Brodmann's area 44), left dorsolateral prefrontal area, left precentral gyrus, left insula, and medial frontal gyrus were also observed. In the basal ganglia, activation in the left thalamus (ventral posteromedial nucleus) and putamen was found. Our results suggest that the primary and secondary somatosensory areas are recruited during tactile imagery, and have partially overlapping neural substrates for the perception of tactile stimulation.     

Speed of lexical decision correlates with diffusion anisotropy in left parietal and frontal white matter: evidence from diffusion tensor imaging

Speed of visual word recognition is an important variable affecting linguistic competence. Although speed of visual word recognition varies widely between individuals, the neural basis of reaction time (RT) differences is poorly understood. Recently, a magnetic resonance technique called diffusion tensor imaging (DTI) has been shown to provide information about white matter (WM) microstructure in vivo. Here, we used DTI to explore whether visual word recognition RT correlates with regional fractional anisotropy (FA) values in the WM of healthy young adults. Participants completed a speeded lexical decision task that involved visual input, linguistic processes, and a motor response output. Results indicated that lexical decision RT was correlated negatively with FA in WM of inferior parietal and frontal language regions rather than in WM of visual or motor regions. Voxels within the inferior parietal and frontal correlation clusters were composed primarily of DTI-based tracts oriented in the anterior-posterior orientation at or near the superior longitudinal fasciculus (SLF) and likely including other smaller association fibers. These results provide new microstructural evidence demonstrating that speed of lexical decision is associated with the degree to which portions of frontal and parietal WM are directionally oriented.     

Attending to and remembering tactile stimuli: a review of brain imaging data and single-neuron responses.

Clinical and neuroimaging observations of the cortical network implicated in tactile attention have identified foci in parietal somatosensory, posterior parietal, and superior frontal locations. Tasks involving intentional hand-arm movements activate similar or nearby parietal and frontal foci. Visual spatial attention tasks and deliberate visuomotor behavior also activate overlapping posterior parietal and frontal foci. Studies in the visual and somatosensory systems thus support a proposal that attention to the spatial location of an object engages cortical regions responsible for the same coordinate referents used for guiding purposeful motor behavior. Tactile attention also biases processing in the somatosensory cortex through amplification of responses to relevant features of selected stimuli. Psychophysical studies demonstrate retention gradients for tactile stimuli like those reported for visual and auditory stimuli, and suggest analogous neural mechanisms for working memory across modalities. Neuroimaging studies in humans using memory tasks, and anatomic studies in monkeys support the idea that tactile information relayed from the somatosensory cortex is directed ventrally through the insula to the frontal cortex for short-term retention and to structures of the medial temporal lobe for long-term encoding. At the level of single neurons, tactile (such as visual and auditory) short-term memory appears as a persistent response during delay intervals between sampled stimuli.     

Perceptual specificity in visual object priming: functional magnetic resonance imaging evidence for a laterality difference in fusiform cortex

Seeing an object on one occasion may facilitate or prime processing of the same object if it is later again encountered. Such priming may also be found -- but at a reduced level -- for different but perceptually similar objects that are alternative exemplars or 'tokens' of the initially presented object. We explored the neural correlates of this perceptual specificity using event-related functional magnetic resonance imaging (fMRI) procedures, contrasting neural activity when participants made object classification decisions (size judgments) regarding previously presented objects (repeated same), alternative exemplars of previously presented objects (repeated different), or entirely new objects (novel). Many frontal regions (including bilateral frontal operculum, bilateral posterior inferior frontal/precentral, left anterior inferior frontal, and superior frontal cortices) and multiple late visual and posterior regions (including middle occipital, fusiform, fusiform-parahippocampal, precuneus, and posterior cingulate, all bilaterally), demonstrated reduced neural activity for repeated compared to novel objects. Greater repetition-induced reductions were observed for same than for different exemplars in several of these regions (bilateral posterior inferior frontal, right precuneus, bilateral middle occipital, bilateral fusiform, bilateral parahippocampal and bilateral superior parietal). Additionally, right fusiform (occipitotemporal) cortex showed significantly less priming for different versus same exemplars than did left fusiform. These findings converge with behavioral evidence from divided visual field studies and with neuropsychological evidence underscoring the key role of right occipitotemporal cortex in processing specific visual form information; possible differences in the representational-functional role of left fusiform are discussed.     

Functional MRI assessment of orofacial articulators: neural correlates of lip, jaw, larynx, and tongue movements

Compared with complex coordinated orofacial actions, few neuroimaging studies have attempted to determine the shared and distinct neural substrates of supralaryngeal and laryngeal articulatory movements when performed independently. To determine cortical and subcortical regions associated with supralaryngeal motor control, participants produced lip, tongue and jaw movements while undergoing functional magnetic resonance imaging (fMRI). For laryngeal motor activity, participants produced the steady-state/i/vowel. A sparse temporal sampling acquisition method was used to minimize movement-related artifacts. Three main findings were observed. First, the four tasks activated a set of largely overlapping, common brain areas: the sensorimotor and premotor cortices, the right inferior frontal gyrus, the supplementary motor area, the left parietal operculum and the adjacent inferior parietal lobule, the basal ganglia and the cerebellum. Second, differences between tasks were restricted to the bilateral auditory cortices and to the left ventrolateral sensorimotor cortex, with greater signal intensity for vowel vocalization. Finally, a dorso-ventral somatotopic organization of lip, jaw, vocalic/laryngeal, and tongue movements was observed within the primary motor and somatosensory cortices using individual region-of-interest (ROI) analyses. These results provide evidence for a core neural network involved in laryngeal and supralaryngeal motor control and further refine the sensorimotor somatotopic organization of orofacial articulators.     

Dual-task interference during initial learning of a new motor task results from competition for the same brain areas

Cerebral patterns of activity elicited by dual-task performance throughout the learning of a complex bimanual coordination pattern were addressed. Subjects (N = 12) were trained on the coordination pattern and scanned using fMRI at early (PRE) and late (POST) learning stages. During scanning, the coordination pattern was performed either as a single task or in concurrence with a simultaneous visual search task (i.e. dual task). Kinematics data revealed a significant performance improvement as a result of learning. In PRE-scanning, the dual-task condition induced deterioration of motor performance, relative to the single-task condition. Activity in lateral frontal and parietal regions involved in both visual search and motor coordination tasks (i.e. ‘overlapping’ regions) was reduced when the tasks were performed simultaneously. In POST-scanning, kinematics performance was equivalent under single- and dual-task conditions, suggesting automaticity of the coordination pattern. Furthermore, overlap between regions involved in visual search and motor tasks was reduced, and dual-task performance was no longer associated with reduction of frontal and parietal activity. Our results suggest that behavioral interference between a complex motor coordination task and a simple simultaneous visual search task occurs when both tasks recruit overlapping regions in the frontal and parietal cortices. This may provide a neural basis for dissipation of dual-task interference following extensive motor practice, which is a traditional behavioral marker of motor automaticity.     

Insular cortex and neuropsychiatric disorders: a review of recent literature.

The insular cortex is located in the centre of the cerebral hemisphere, having connections with the primary and secondary somatosensory areas, anterior cingulate cortex, amygdaloid body, prefrontal cortex, superior temporal gyrus, temporal pole, orbitofrontal cortex, frontal and parietal opercula, primary and association auditory cortices, visual association cortex, olfactory bulb, hippocampus, entorhinal cortex, and motor cortex. Accordingly, dense connections exist among insular cortex neurons. The insular cortex is involved in the processing of visceral sensory, visceral motor, vestibular, attention, pain, emotion, verbal, motor information, inputs related to music and eating, in addition to gustatory, olfactory, visual, auditory, and tactile data. In this article, the literature on the relationship between the insular cortex and neuropsychiatric disorders was summarized following a computer search of the Pub-Med database. Recent neuroimaging data, including voxel based morphometry, PET and fMRI, revealed that the insular cortex was involved in various neuropsychiatric diseases such as mood disorders, panic disorders, PTSD, obsessive-compulsive disorders, eating disorders, and schizophrenia. Investigations of functions and connections of the insular cortex suggest that sensory information including gustatory, olfactory, visual, auditory, and tactile inputs converge on the insular cortex, and that these multimodal sensory information may be integrated there.     

The neural substrate of gesture recognition

Previous studies have linked action recognition with a particular pool of neurons located in the ventral premotor cortex, the posterior parietal cortex and the superior temporal sulcus (the mirror neuron system). However, it is still unclear if transitive and intransitive gestures share the same neural substrates during action-recognition processes. In the present study, we used event-related functional magnetic resonance imaging (fMRI) to assess the cortical areas active during recognition of pantomimed transitive actions, intransitive gestures, and meaningless control actions. Perception of all types of gestures engaged the right pre-supplementary motor area (pre-SMA), and bilaterally in the posterior superior temporal cortex, the posterior parietal cortex, occipitotemporal regions and visual cortices. Activation of the posterior superior temporal sulcus/superior temporal gyrus region was found in both hemispheres during recognition of transitive and intransitive gestures, and in the right hemisphere during the control condition; the middle temporal gyrus showed activation in the left hemisphere when subjects recognized transitive and intransitive gestures; activation of the left inferior parietal lobe and intraparietal sulcus (IPS) was mainly observed in the left hemisphere during recognition of the three conditions. The most striking finding was the greater activation of the left inferior frontal gyrus (IFG) during recognition of intransitive actions. Results show that a similar neural substrate, albeit, with a distinct engagement underlies the cognitive processing of transitive and intransitive gestures recognition. These findings suggest that selective disruptions in these circuits may lead to distinct clinical deficits.     

Sensory‐motor brain network connectivity for speech comprehension

The act of listening to speech activates a large network of brain areas. In the present work, a novel data‐driven technique (the combination of independent component analysis and Granger causality) was used to extract brain network dynamics from an fMRI study of passive listening to Words, Pseudo‐Words, and Reverse‐played words. Using this method we show the functional connectivity modulations among classical language regions (Broca's and Wernicke's areas) and inferior parietal, somatosensory, and motor areas and right cerebellum. Word listening elicited a compact pattern of connectivity within a parieto‐somato‐motor network and between the superior temporal and inferior frontal gyri. Pseudo‐Word stimuli induced activities similar to the Word condition, which were characterized by a highly recurrent connectivity pattern, mostly driven by the temporal lobe activity. Also the Reversed‐Word condition revealed an important influence of temporal cortices, but no integrated activity of the parieto‐somato‐motor network. In parallel, the right cerebellum lost its functional connection with motor areas, present in both Word and Pseudo‐Word listening. The inability of the participant to produce the Reversed‐Word stimuli also evidenced two separate networks: the first was driven by frontal areas and the right cerebellum toward somatosensory cortices; the second was triggered by temporal and parietal sites towards motor areas. Summing up, our results suggest that semantic content modulates the general compactness of network dynamics as well as the balance between frontal and temporal language areas in driving those dynamics. The degree of reproducibility of auditory speech material modulates the connectivity pattern within and toward somatosensory and motor areas. Hum Brain Mapp, 2010. © 2009 Wiley‐Liss, Inc.     

Shared and dissociated cortical regions for object and letter processing

The present study determined the extent to which object and letter recognition recruit similar or dissociated neural resources. Participants passively viewed and silently named line drawings of objects, single letters, and visual noise patterns and centrally fixated an asterisk. We used whole-brain functional MRI and a very conservative approach to hypothesis testing that distinguished among brain regions that were selectively activated by different experimental conditions and those that were conjointly activated. The left fusiform gyrus (BA 19 & 37) and left inferior frontal cortex BA(44/6) showed a greater degree of conjoined activation for objects and letters than selective activation for either category, whereas left inferior parietal cortex (BA 40) and the left insula showed a strong letter-selective response. Equal recruitment of left fusiform and inferior frontal regions by objects and letters reflects similar demands on cognitive processing by these two categories and argues against category-specific modules in these regions. However, cortical systems for object and letter processing are not completely shared given the exclusive activation of left inferior parietal cortex by letters.     

Neural substrates for observing and imagining non-object-directed actions

The present fMRI study was aimed at assessing the cortical areas active when individuals observe non-object-directed actions (mimed, symbolic, and meaningless), and when they imagine performing those same actions. fMRI signal increases in common between action observation and motor imagery were found in the premotor cortex and in a large region of the inferior parietal lobule. While the premotor cortex activation overlapped that previously found during the observation and imagination of object-directed actions, in the parietal lobe the signal increase was not restricted to the intraparietal sulcus region, known to be active during the observation and imagination of object-directed actions, but extended into the supramarginal and angular gyri. When contrasting motor imagery with the observation of non-object-directed actions, signal increases were found in the mesial frontal and cingulate cortices, the supramarginal gyrus, and the inferior frontal gyrus. The opposite contrast showed activation virtually limited to visual areas. In conclusion, the present data define the common circuit for observing and imagining non-object-directed actions. In addition, they show that the representation of non-object-directed actions include parietal regions not found to be involved in coding object-directed actions.     

Role of the corpus callosum in the somatosensory activation of the ipsilateral cerebral cortex: an fMRI study of callosotomized patients

To verify whether the activation of the posterior parietal and parietal opercular cortices to tactile stimulation of the ipsilateral hand is mediated by the corpus callosum, a functional magnetic resonance imaging (fMRI, 1.0 tesla) study was performed in 12 control and 12 callosotomized subjects (three with total and nine with partial resection). Eleven patients were also submitted to the tactile naming test. In all subjects, unilateral tactile stimulation provoked a signal increase temporally correlated with the stimulus in three cortical regions of the contralateral hemisphere. One corresponded to the first somatosensory area, the second was in the posterior parietal cortex, and the third in the parietal opercular cortex. In controls, activation was also observed in the ipsilateral posterior parietal and parietal opercular cortices, in regions anatomically corresponding to those activated contralaterally. In callosotomized subjects, activation in the ipsilateral hemisphere was observed only in two patients with splenium and posterior body intact. These two patients and another four with the entire splenium and variable portions of the posterior body unsectioned named objects explored with the right and left hand without errors. This ability was impaired in the other patients. The present physiological and anatomical data indicate that in humans activation of the posterior parietal and parietal opercular cortices in the hemisphere ipsilateral to the stimulated hand is mediated by the corpus callosum, and that the commissural fibres involved probably cross the midline in the posterior third of its body.     

Perceptual and Semantic Representations at Encoding Contribute to True and False Recognition of Objects

When encoding new episodic memories, visual and semantic processing is proposed to make distinct contributions to accurate memory and memory distortions. Here, we used fMRI and preregistered representational similarity analysis to uncover the representations that predict true and false recognition of unfamiliar objects. Two semantic models captured coarse-grained taxonomic categories and specific object features, respectively, while two perceptual models embodied low-level visual properties. Twenty-eight female and male participants encoded images of objects during fMRI scanning, and later had to discriminate studied objects from similar lures and novel objects in a recognition memory test. Both perceptual and semantic models predicted true memory. When studied objects were later identified correctly, neural patterns corresponded to low-level visual representations of these object images in the early visual cortex, lingual, and fusiform gyri. In a similar fashion, alignment of neural patterns with fine-grained semantic feature representations in the fusiform gyrus also predicted true recognition. However, emphasis on coarser taxonomic representations predicted forgetting more anteriorly in the anterior ventral temporal cortex, left inferior frontal gyrus and, in an exploratory analysis, left perirhinal cortex. In contrast, false recognition of similar lure objects was associated with weaker visual analysis posteriorly in early visual and left occipitotemporal cortex. The results implicate multiple perceptual and semantic representations in successful memory encoding and suggest that fine-grained semantic as well as visual analysis contributes to accurate later recognition, while processing visual image detail is critical for avoiding false recognition errors.SIGNIFICANCE STATEMENT People are able to store detailed memories of many similar objects. We offer new insights into the encoding of these specific memories by combining fMRI with explicit models of how image properties and object knowledge are represented in the brain. When people processed fine-grained visual properties in occipital and posterior temporal cortex, they were more likely to recognize the objects later and less likely to falsely recognize similar objects. In contrast, while object-specific feature representations in fusiform gyrus predicted accurate memory, coarse-grained categorical representations in frontal and temporal regions predicted forgetting. The data provide the first direct tests of theoretical assumptions about encoding true and false memories, suggesting that semantic representations contribute to specific memories as well as errors.     

Executive functions with different motor outputs in somatosensory Go/Nogo tasks: an event-related functional MRI study

The aim of this event-related functional magnetic resonance imaging (fMRI) study was to investigate and compare executive functions with different motor outputs in somatosensory Go/Nogo tasks: (1) Button press and (2) Count. Go and Nogo stimuli were presented with an even probability. We observed a common network for Movement and Count Go trials in several regions of the brain including the dorsolateral (DLPFC) and ventrolateral prefrontal cortices (VLPFC), supplementary motor area (SMA), posterior parietal cortex (PPC), inferior parietal lobule (IPL), Insula, and superior temporal gyrus (STG). Direct comparison revealed that primary sensorimotor area (SMI), premotor area (PM), and anterior cingulate cortex (ACC) were more activated during Movement than Count Go trials. In contrast, the VLPFC was more activated during Count than Movement Go trials. Our results suggest that there were two neural networks for the supramodal executive function, common and uncommon, depending on the required response mode.     

An event-related functional MRI study comparing interference effects in the Simon and Stroop tasks

The Stroop and Simon tasks typify a class of interference effects in which the introduction of task-irrelevant stimulus characteristics robustly slows reaction times. Behavioral studies have not succeeded in determining whether the neural basis for the resolution of these interference effects during successful task performance is similar or different across tasks. Event-related functional magnetic resonance imaging (fMRI) studies were obtained in 10 healthy young adults during performance of the Stroop and Simon tasks. Activation during the Stroop task replicated findings from two earlier fMRI studies. These activations were remarkably similar to those observed during the Simon task, and included anterior cingulate, supplementary motor, visual association, inferior temporal, inferior parietal, inferior frontal, and dorsolateral prefrontal cortices, as well as the caudate nuclei. The time courses of activation were also similar across tasks. Resolution of interference effects in the Simon and Stroop tasks engage similar brain regions, and with a similar time course. Therefore, despite the widely differing stimulus characteristics employed by these tasks, the neural systems that subserve successful task performance are likely to be similar as well.     

Object priming and recognition memory: Dissociable effects in left frontal cortex at encoding

Functional magnetic resonance imaging (fMRI) studies have implicated the left prefrontal cortex in priming. We tested the hypothesis that object encoding activity in different prefrontal cortex regions selectively predicts subsequent object priming and recognition respectively. Participants were scanned whilst making semantic category judgements about novel object pictures. One week later priming and recognition of these objects were tested. Encoding that produced long-lasting priming in the absence of recognition memory was associated with increased activity in left inferior prefrontal (BA 47) and superior frontal (BA 8) cortices. In contrast, encoding that produced object recognition one week later activated the left middle frontal cortex (BA 9). This is consistent with other evidence indicating that object priming and recognition are independent kind of memory. Problems of measuring item-by-item recognition and priming together are discussed.     

Remembrance of things touched: how sensorimotor experience affects the neural instantiation of object form

Numerous neuroimaging and neuropsychological studies have highlighted the role of the ventral, occipitotemporal visual processing stream in the representation and retrieval of semantic memory for the appearance of objects. Here, we examine the role of the parietal cortex in retrieval of object shape information. fMRI data were acquired as subjects listened to the names of common objects and made judgments about their shape. Recruitment of the left inferior parietal lobule (IPL) during shape retrieval was modulated by the amount of prior tactile experience associated with the objects. In addition, the same pattern of activity was observed in right postcentral gyrus, suggesting that the representation of object shape is distributed amongst regions that are relevant to the sensorimotor acquisition history of this attribute, as predicted by distributed models of semantic memory.     

Communication with emblematic gestures: shared and distinct neural correlates of expression and reception

Emblematic (or symbolic) gestures allow individuals to convey a variety of thoughts and emotions ranging from approval to hostility. The use of such gestures involves the execution of a codified motor act by the addresser and its perception and decoding by the addressee. To examine underlying common and distinct neural correlates, we used fMRI tasks in which subjects viewed video clips of emblematic one-hand gestures. They were asked to (1) take the perspective of the addresser and imagine executing the gestures ("expression" condition), and to (2) take the perspective of the addressee and imagine being confronted with the gestures ("reception" condition). Common areas of activation were found in inferior frontal, medial frontal, and posterior temporal cortices with left-hemispheric predominance as well as in the cerebellum. The distinction between regions specifically involved in the expression or reception condition partly resembled the dorsal and ventral stream dichotomy of visual processing with junctions in inferior frontal and medial prefrontal cortices. Imagery of gesture expression involved the dorsal visual stream as well as higher-order motor areas. In contrast, gesture reception encompassed regions related to semantic processing, and medial prefrontal areas known to be involved in the process of understanding the intentions of others. In conclusion, our results provide evidence for a dissociation in representations of emblematic gesture processing between addresser and addressee in addition to shared components in language-related areas.