The sightless view: neural correlates of occluded objects

The relationship between neural activity and object perception has received considerable attention using stimulus manipulations such as masking or dichoptic presentation. Here we investigate the same problem by occluding objects with an opaque screen that acts to dissociate the direct perception of the object from the awareness of its presence. We used functional magnetic resonance imaging to measure brain activity when subjects viewed objects (faces and houses) that underwent occlusion and found that the response of the majority of the fusiform face area (FFA) and lateral occipital cortex is the same whether the object is visible or occluded. This suggests that when objects are directly viewed, activity within object-selective regions may reflect the awareness of presence, not the direct perception, of the object. Additionally, we identify a region of premotor cortex that is selectively activated by occlusion of either object type, suggesting its generic involvement with processing occluded objects.     

Explicit neural representations,recursive neural networks and conscious visual perception

The fundamental question as to whether the neural correlates of any given conscious visual experience are expressed locally within a given cortical area or more globally within some widely distributed network remains unresolved. We inquire as to whether recursive processing-by which we mean the combined flow and integrated outcome of afferent and recurrent activity across a series of cortical areas-is essential for the emergence of conscious visual experience. If so, we further inquire as to whether such recursive processing is essential only for loops between extrastriate cortical areas explicitly representing experiences such as color or motion back to V1 or whether it is processing between still higher levels and the areas computing such explicit representations that is exclusively or additionally essential for visual experience. If recursive processing is not essential for the emergence of conscious visual experience, then it should also be possible to determine whether it is only the intracortical sensory processing within areas computing explicit sensory representations that is required for perceptual experience or whether it is the subsequent processing of the output of such areas within more anterior cortical regions that engenders perception. The present analysis suggests that the questions posed here may ultimately become experimentally resolvable. Whatever the outcome, the results will likely open new approaches to identify the neural correlates of conscious visual perception.     

The effects of age on the neural correlates of episodic encoding

Young and old adults underwent positron emission tomographic scans while encoding pictures of objects and words using three encoding strategies: deep processing (a semantic living/nonliving judgement), shallow processing (size judgement) and intentional learning. Picture memory exceeded word memory in both young and old groups, and there was an age-related decrement only in word recognition. During the encoding tasks three brain activity patterns were found that differentiated stimulus type and the different encoding strategies. The stimulus-specific pattern was characterized by greater activity in extrastriate and medial temporal cortices during picture encoding, and greater activity in left prefrontal and temporal cortices during encoding of words. The older adults showed this pattern to a significantly lesser degree. A pattern distinguishing deep processing from intentional learning of words and pictures was identified, characterized mainly by differences in prefrontal cortex, and this pattern also was of significantly lesser magnitude in the old group. A final pattern identified areas with increased activity during deep processing and intentional learning of pictures, including left prefrontal and bilateral medial temporal regions. There was no group difference in this pattern. These results indicate age-related dysfunction in several encoding networks, with sparing of one specifically involved in more elaborate encoding of pictures. These age-related changes appear to affect verbal memory more than picture memory.     

Studies on colonic motor correlates of spontaneous defecation in conscious dogs

The colonic motilities during defecation were studied by means of extraluminal strain gauge force transducers in six conscious dogs. A set of eight transducers was implanted in each dog: one was on the terminal ileum and the remaining seven were on the whole length of the colon equidistantly. As Karaus & Sarna had already described in 1987, giant migrating contractions (GMC) that were initiated in the proximal colon and rapidly migrated caudad before defecation were reconfirmed also in this study to be motor equivalent of mass movements. Several new findings in addition were obtained. At spontaneous defecation, a reflex relaxation was always observed at the distal end of the colon. This colonic outlet relaxation (COR) usually occurred synchronously with the initiation of GMC at the proximal colon and lasted until GMC had arrived at the distal end. Evacuation of feces occurred during this COR. COR was not observed at defecation induced by neostigmine or prostaglandin F2 alpha. After bilateral pelvic nerve section, both GMC and COR were completely abolished. Instead, group of low amplitude caudad migrating contractions occurred at the proximal or middle colon and were followed by frequent evacuation of small amount of loose stools. COR was not observed at this type of defecation. It is suggested from this study that not only GMC but also COR are the essential motor correlates of spontaneous defecation. Both GMC and COR are under control of pelvic nerves.     

The neural correlates of human working memory for haptically explored object orientations

Skillful object manipulation requires that haptically explored spatial object characteristics like orientation be adequately represented in working memory. In the current functional magnetic resonance imaging study, healthy right-handed participants explored a bar-shaped reference object with the left hand, memorizing its orientation. After a variable delay (0.5, 5, or 10 s), participants used their right hand to match the orientation by rotating a second, identical object. In the first seconds of the delay, right sensorimotor cortex was active, whereas clusters in left anterior prefrontal cortex (aPFC) (Brodmann area 10) became dominant 2 s after the end of exploration, showing sustained activity for several seconds. In contrast, left parieto-occipital cortex was involved toward the end of the delay interval. Our results indicate that a dynamic network of brain areas subserves hapticospatial information processing in the delay between haptic stimulus exploration and orientation matching. We propose that haptic sensory traces, maintained in contralateral sensorimotor cortex, are transformed into more abstract hapticospatial representations in the early delay stages. Maintenance of these representations engages aPFC and parieto-occipital cortex. Whereas aPFC possibly integrates spatial and motor components of hapticospatial working memory, parieto-occipital cortex might be involved in orientation imagery, supporting working memory, and the preparation of haptic matching.     

Psychological structure and neural correlates of event knowledge

Humans are capable of storing and retrieving sequences of complex structured events. Here we report a study in which we establish the psychological structure of event knowledge and then use parametric event-related functional magnetic resonance imaging to identify its neural correlates. We demonstrate that event knowledge is organized along dissociable dimensions that are reflected in distinctive patterns of neural activation: social valence (amygdala and right orbitofrontal cortex), experience (medial prefrontal cortex) and engagement (left orbitofrontal cortex). Our study affirms the importance and uniqueness of the human prefrontal cortex in representing event knowledge.     

The effects of aging on the neural correlates of subjective and objective recollection

High-functioning older adults can exhibit normal recollection when measured subjectively, via "remember" judgments, but not when measured objectively, via source judgments, whereas low-functioning older adults exhibit impairments for both measures. A potential explanation for this is that typical subjective and objective tests of recollection necessitate different processing demands, supported by distinct brain regions, and that deficits in these tests are observed according to the degree of age-related changes in these regions. Here, we used event-related functional magnetic resonance imaging to measure the effects of aging on neural correlates of subjective and objective measures of recollection, in young, high-functioning (Old-High) and low-functioning (Old-Low) older adults. Behaviorally, the Old-High group showed intact subjective ("remember" judgments) but impaired objective recollection (for 1 of 2 spatial or temporal sources), whereas the Old-Low group was impaired on both measures. Imaging data showed changes in parietal subjective recollection effects in the Old-Low group and in lateral frontal objective recollection effects in both older adult groups. Our results highlight the importance of examining performance variability in older adults and suggest that differential effects of aging on brain regions are associated with different patterns of performance on tests of subjective and objective recollection.     

Typical neural representations of action verbs develop without vision

Many empiricist theories hold that concepts are composed of sensory-motor primitives. For example, the meaning of the word "run" is in part a visual image of running. If action concepts are partly visual, then the concepts of congenitally blind individuals should be altered in that they lack these visual features. We compared semantic judgments and neural activity during action verb comprehension in congenitally blind and sighted individuals. Participants made similarity judgments about pairs of nouns and verbs that varied in the visual motion they conveyed. Blind adults showed the same pattern of similarity judgments as sighted adults. We identified the left middle temporal gyrus (lMTG) brain region that putatively stores visual-motion features relevant to action verbs. The functional profile and location of this region was identical in sighted and congenitally blind individuals. Furthermore, the lMTG was more active for all verbs than nouns, irrespective of visual-motion features. We conclude that the lMTG contains abstract representations of verb meanings rather than visual-motion images. Our data suggest that conceptual brain regions are not altered by the sensory modality of learning.     

Dissociating neural correlates of cognitive components in mental calculation

Mental calculation is a complex cognitive operation that is composed of a set of distinct functional processes. Using functional magnetic resonance imaging (fMRI), we mapped brain activity in healthy subjects performing arithmetical tasks and control tasks evoking a comparable load on visuo-constructive, linguistic, attentional and mnemonic functions. During calculation, as well as non-mathematical tasks, similar cortical networks consisting of bilateral prefrontal, premotor and parietal regions were activated, suggesting that most of these cortical areas do not exclusively represent modules for calculation but support more general cognitive operations that are instrumental but not specific to mental arithmetic. Significant differences between calculation and the non-mathematical tasks were found in parietal sub-regions, where non-arithmetic number or letter substitution tasks preferentially activated the superior parietal lobules whereas calculation predominantly elicited activation of the left dorsal angular gyrus and the medial parietal cortices. We interpret the latter activations to reflect sub-processes of mental calculation that are related to the processing of numerical representations during exact calculation and to arithmetical fact retrieval. Finally, we found that more complex calculation tasks involving the application of calculation rules increased activity in left inferior frontal areas that are known to subserve linguistic and working memory functions. Taken together, these findings help to embed the specific cognitive operation of calculation into a neural framework that provides the required set of instrumental components. This result may further inform the cognitive modeling of calculation and adds to the understanding of neuropsychological deficit patterns in patients.     

The effects of aging on material-independent and material-dependent neural correlates of source memory retrieval

Age-related declines in source memory have been observed for various stimuli and associated details. These impairments may be related to alterations in brain regions contributing to source memory via material-independent processes and/or regions specialized for processing specific materials. Using event-related functional magnetic resonance imaging, we investigate the effects of aging on source memory and associated neural activity for words and objects. Source accuracy was equally impaired in older adults for both materials. Imaging data revealed both groups recruited similar networks of regions to support source memory accuracy irrespective of material, including parietal and prefrontal cortices (PFC) and the hippocampus. Age-related decreases in material-independent activity linked to postretrieval monitoring were observed in right lateral PFC. Additionally, age-related increases in source accuracy effects were shown in perirhinal cortex, which were positively correlated with performance in older adults, potentially reflecting functional compensation. In addition to group differences in material-independent regions, age-related crossover interactions for material-dependent source memory effects were observed in regions selectively engaged by objects. These results suggest that older adults' source memory impairments reflect alterations in regions making material-independent contributions to source memory retrieval, primarily the lateral PFC, but may be further impacted by changes in regions sensitive to particular materials.     

The relationship between aging, performance, and the neural correlates of successful memory encoding

The present functional magnetic resonance imaging (fMRI) study investigated whether age-related differences in the neural correlates of successful memory encoding are modulated by memory performance. Young (mean age 22 years; N = 16) and older (mean age 69 years; N = 32) subjects were scanned while making animacy decisions on visually presented words. Memory for the words was later assessed in a recognition test, allowing fMRI activity elicited by study words to be contrasted according to subsequent memory performance. Young and older adults exhibited equivalent subsequent memory effects (enhanced activity for later remembered items) in an extensive network that included left inferior prefrontal cortex and anterior hippocampus. In posterior cingulate cortex, reversed subsequent memory effects (greater activity for later forgotten items) were of greater magnitude in young subjects. A voxel-of-interest analysis conducted on left and right prefrontal subsequent memory effects revealed that the effects were distributed more bilaterally in older than in young subjects, replicating previous findings. This age-related difference was confined to older subjects with relatively poor recognition performance, who were also the only group to demonstrate statistically significant right prefrontal subsequent memory effects. The findings suggest that relative preservation of memory performance with increasing age does not depend upon right prefrontal "over-recruitment."     

Biliary sphincter motility after neural isolation of the pancreatoduodenal region in conscious dogs

BACKGROUND: Several neural and hormonal factors are known to affect the motility of the sphincter of Oddi. However, the precise mechanisms of the control of sphincter motility have not been completely explored. We investigated the relationship of canine biliary sphincter motility when it is extrinsically denervated by neural isolation of the pancreatoduodenal region. METHODS: Interdigestive and postprandial sphincter motility in a denervated pancreatoduodenal segment and effects of cholecystokinin-octapeptide were studied in 7 conscious dogs. Data were compared with those of 7 neurally intact control dogs. RESULTS: After extrinsic denervation of the pancreatoduodenal region, sphincter motility exerted a cyclic change in concert with the duodenal myoelectric cycles; this change involved short cyclic bursts of motor activity, which gradually increased in intensity. The increase in the cyclic bursts of motor activity was also cyclic and associated with an increase in the plasma motilin concentration. Neural isolation of the pancreatoduodenal region increased sphincter basal pressure and motility index (integral per minute). In the denervated biliary sphincter, the feeding pattern and temporary inhibitory effect of feeding, as seen in controls, were absent, which suggests the role of extrinsic nerves in delivering bile into the duodenum after feeding. In the denervated dogs, cholecystokinin-octapeptide caused excitation of the sphincter activity, instead of relaxation observed in controls. CONCLUSIONS: Extrinsic innervation to the pancreatoduodenal region has an inhibitory effect on biliary sphincter motility. Abnormalities in extrinsic innervation to the biliary sphincter might increase the resistance of the sphincter to the bile flow and induce bile stagnation.     

Water intake and the neural correlates of the consciousness of thirst

Thirst and resultant water drinking can arise in response to deficits in both the intracellular and extracellular fluid compartments. Inhibitory influences mediating the satiation of thirst also are necessary to prevent overhydration. The brain regions that underpin the generation or inhibition of thirst in these circumstances can be categorized as sensory, integrative, or cortical effector sites. The anterior cingulate cortex and insula are activated in thirsty human beings as shown by functional brain-imaging techniques. It is postulated that these sites may be cortical effector regions for thirst. A major sensory site for generating thirst is the lamina terminalis in the forebrain. Osmoreceptors within the organum vasculosum of the lamina terminalis and subfornical organ detect systemic hypertonicity. The subfornical organ mediates the dipsogenic actions of circulating angiotensin II and relaxin. Major integrative sites are the nucleus of the tractus solitarius, the lateral parabrachial nucleus, the midbrain raphé nuclei, the median preoptic nucleus, and the septum. Despite these advances, most of the neural pathways and neurochemical mechanisms subserving the genesis of thirst remain to be elucidated.     

The neural correlates of declining performance with age: evidence for age-related changes in cognitive control

The neural system involved in cognitive control includes the anterior cingulate cortex (ACC) and the lateral prefrontal cortex (PFC). Neural activity within these structures is sensitive to aging. We investigated the hypothesis that decline in performance with age results in increased cognitive control, as indexed by greater activity within the ACC and lateral PFC. Using positron emission tomography we measured neural activity during a range of verbal decision-making tasks in 16 subjects aged 37-83 years. Conditions were separated behaviorally on the basis of their sensitivity to aging. This allowed the comparison of age-dependent and age-independent conditions, revealing the neural correlates of age-dependent decline in performance. We then modeled the relationship between age, decision type, performance, and frontal lobe activity. ACC activity was independently predicted by age and decision-making accuracy, indicating that in older individuals ACC response is more sensitive to declining performance. We also found strong functional connectivity between the ACC and lateral PFC and observed that activation of the lateral PFC was qualitatively different over time in different age groups. Thus, the ACC and lateral PFC show distinct responses to age-related decline in decision-making performance. This suggests that greater cognitive control is employed as individuals age and their performance declines.     

Decomposing the neural correlates of antisaccade eye movements using event-related FMRI

The antisaccade task is a model of the conflict between an unwantedreflexive response (which must be inhibited) and a complex volitionalresponse (which must be generated). The present experiment aimedto investigate separately the neural correlates of these cognitivecomponents using a delayed saccade paradigm to dissociate saccadeinhibition from generation. Seventeen healthy volunteers completedevent-related functional magnetic resonance imaging at 1.5 Tduring saccades to and away from a peripheral visual target(prosaccades and antisaccades, respectively). Saccades wererequested in response to an auditory go signal on average 12s after peripheral target appearance. It was found that theright supramarginal gyrus showed significantly greater activationduring the inhibition phase than the generation phase of theparadigm for both antisaccade and prosaccade trials, suggestinga role in saccade inhibition or stimulus detection. On the otherhand, the right lateral frontal eye field and bilateral intraparietalsulcus showed evidence of selective involvement in antisaccadegeneration. Ventrolateral and dorsolateral prefrontal corticesshowed comparable levels of activation in both phases of thetask. These areas likely fulfill a more general supervisoryrole in the volitional control of eye movements, such as stimulusappraisal, task set, and decision making.     

The different neural correlates of action and functional knowledge in semantic memory: an FMRI study

Previous reports suggest that the internal organization of semantic memory is in terms of different "types of knowledge," including "sensory" (information about perceptual features), "action" (motor-based knowledge of object utilization), and "functional" (abstract properties, as function and context of use). Consistent with this view, a specific loss of action knowledge, with preserved functional knowledge, has been recently observed in patients with left frontoparietal lesions. The opposite pattern (impaired functional knowledge with preserved action knowledge) was reported in association with anterior inferotemporal lesions. In the present study, the cerebral representation of action and functional knowledge was investigated using event-related analysis of functional magnetic resonance imaging data. Fifteen subjects were presented with pictures showing pairs of manipulable objects and asked whether the objects within each pair were used with the same manipulation pattern ("action knowledge" condition) or in the same context ("functional knowledge" condition). Direct comparisons showed action knowledge, relative to functional knowledge, to activate a left frontoparietal network, comprising the intraparietal sulcus, the inferior parietal lobule, and the dorsal premotor cortex. The reverse comparison yielded activations in the retrosplenial and the lateral anterior inferotemporal cortex. These results confirm and extend previous neuropsychological data and support the hypothesis of the existence of different types of information processing in the internal organization of semantic memory.     

The spatiotemporal dynamics of autobiographical memory: neural correlates of recall, emotional intensity, and reliving

We sought to map the time course of autobiographical memory retrieval, including brain regions that mediate phenomenological experiences of reliving and emotional intensity. Participants recalled personal memories to auditory word cues during event-related functional magnetic resonance imaging (fMRI). Participants pressed a button when a memory was accessed, maintained and elaborated the memory, and then gave subjective ratings of emotion and reliving. A novel fMRI approach based on timing differences capitalized on the protracted reconstructive process of autobiographical memory to segregate brain areas contributing to initial access and later elaboration and maintenance of episodic memories. The initial period engaged hippocampal, retrosplenial, and medial and right prefrontal activity, whereas the later period recruited visual, precuneus, and left prefrontal activity. Emotional intensity ratings were correlated with activity in several regions, including the amygdala and the hippocampus during the initial period. Reliving ratings were correlated with activity in visual cortex and ventromedial and inferior prefrontal regions during the later period. Frontopolar cortex was the only brain region sensitive to emotional intensity across both periods. Results were confirmed by time-locked averages of the fMRI signal. The findings indicate dynamic recruitment of emotion-, memory-, and sensory-related brain regions during remembering and their dissociable contributions to phenomenological features of the memories.     

Controlling emotional expression: behavioral and neural correlates of nonimitative emotional responses

Emotional facial expressions can engender similar expressions in others. However, adaptive social and motivational behavior can require individuals to suppress, conceal, or override prepotent imitative responses. We predicted, in line with a theory of "emotion contagion," that when viewing a facial expression, expressing a different emotion would manifest as behavioral conflict and interference. We employed facial electromyography (EMG) and functional magnetic resonance imaging (fMRI) to investigate brain activity related to this emotion expression interference (EEI) effect, where the expressed response was either concordant or discordant with the observed emotion. The Simon task was included as a nonemotional comparison for the fMRI study. Facilitation and interference effects were observed in the latency of facial EMG responses. Neuroimaging revealed activation of distributed brain regions including anterior right inferior frontal gyrus (brain area [BA] 47), supplementary motor area (facial area), posterior superior temporal sulcus (STS), and right anterior insula during emotion expression-associated interference. In contrast, nonemotional response conflict (Simon task) engaged a distinct frontostriatal network. Individual differences in empathy and emotion regulatory tendency predicted the magnitude of EEI-evoked regional activity with BA 47 and STS. Our findings point to these regions as providing a putative neural substrate underpinning a crucial adaptive aspect of social/emotional behavior.