Showing papers in "Journal of Cognitive Neuroscience in 1999"
TL;DR: The results indicate that multisensory integration is mediated by flexible, highly adaptive physiological processes that can take place very early in the sensory processing chain and operate in both sensory-specific and nonspecific cortical structures in different ways.
Abstract: The aim of this study was (1) to provide behavioral evidence for multimodal feature integration in an object recognition task in humans and (2) to characterize the processing stages and the neural structures where multisensory interactions take place. Event-related potentials (ERPs) were recorded from 30 scalp electrodes while subjects performed a forcedchoice reaction-time categorization task: At each trial, the subjects had to indicate which of two objects was presented by pressing one of two keys. The two objects were defined by auditory features alone, visual features alone, or the combination of auditory and visual features. Subjects were more accurate and rapid at identifying multimodal than unimodal objects. Spatiotemporal analysis of ERPs and scalp current densities revealed several auditory-visual interaction components temporally, spatially, and functionally distinct before 200 msec poststimulus. The effects observed were (1) in visual areas, new neural activities (as early as 40 msec poststimulus) and modu lation (amplitude decrease) of the N185 wave to unimodal visual stimulus, (2) in the auditory cortex, modulation (amplitude increase) of subcomponents of the unimodal auditory N1 wave around 90 to 110 msec, and (3) new neural activity over the right fronto-temporal area (140 to 165 msec).Furthermore, when the subjects were separated into two groups according to their dominant modality to perform the task in unimodal conditions (shortest reaction time criteria), the integration effects were found to be similar for the two groups over the nonspecific fronto-temporal areas, but they clearly differed in the sensory-specific cortices, affecting predominantly the sensory areas of the nondominant modality. Taken together, the results indicate that multisensory integration is mediated by flexible, highly adaptive physiological processes that can take place very early in the sensory processing chain and operate in both sensory-specific and nonspecific cortical structures in different ways.
1,061 citations
TL;DR: A model that proposes that perceptual tasks interrupt processes ongoing during rest that involve many of the same brain areas engaged during semantic retrieval may help to explain several unanticipated results from prior studies of semantic processing.
Abstract: Localized, task-induced decreases in cerebral blood flow are a frequent finding in functional brain imaging research but remain poorly understood. One account of these phenomena postulates processes ongoing during conscious, resting states that are interrupted or inhibited by task performance. Psychological evidence suggests that conscious humans are engaged almost continuously in adaptive processes involving semantic knowledge retrieval, representation in awareness, and directed manipulation of represented knowledge for organization,problem-solving, and planning. If interruption of such ''conceptual'' processes accounts for task-induced deactivation, tasks that also engage these conceptual processes should not cause deactivation. Furthermore, comparisons between conceptual and nonconceptual tasks should show activation during conceptual tasks of the same brain areas that are ''deactivated'' relative to rest. To test this model, functional magnetic resonance imaging data were acquired during a resting state,a perceptual task,and a semantic retrieval task. A network of left-hemisphere polymodal cortical regions showed higher signal values during the resting state than during the perceptual task but equal values during the resting and semantic conditions. This result is consistent with the proposal that perceptual tasks interrupt processes ongoing during rest that involve many of the same brain areas engaged during semantic retrieval. As further evidence for this model, the same network of brain areas was activated in two direct comparisons between semantic and perceptual processing tasks. This same ''conceptual processing'' network was also identified in several previous studies that contrasted semantic and perceptual tasks or resting and active states. The model proposed here offers a unified account of these findings and may help to explain several unanticipated results from prior studies of semantic processing.
1,037 citations
TL;DR: The results of cortical activity support the hypothesis that motor imagery and motor performance possess similar neural substrates as well as the assumption that the posterior cerebellum is involved in the inhibition of movement execution during imagination.
Abstract: Brain activation during executed (EM) and imagined movements (IM) of the right and left hand was studied in 10 healthy right-handed subjects using functional magnetic resonance imagining (fMRI). Low electromyographic (EMG) activity of the musculi flexor digitorum superficialis and high vividness of the imagined movements were trained prior to image acquisition. Regional cerebral activation was measured by fMRI during EM and IM and compared to resting conditions. Anatomically selected regions of interest (ROIs) were marked interactively over the entire brain. In each ROI activated pixels above a t value of 2.45 (p < 0.01) were counted and analyzed. In all subjects the supplementary motor area (SMA), the premotor cortex (PMC), and the primary motor cortex (M1) showed significant activation during both EM and IM; the somatosen sory cortex (S1) was significantly activated only during EM. Ipsilateral cerebellar activation was decreased during IM compared to EM. In the cerebellum, IM and EM differed in their foci of maximal activation: Highest ipsilateral activation of the cerebellum was observed in the anterior lobe (Larsell lobule H IV) during EM, whereas a lower maximum was found about 2-cm dorsolateral (Larsell lobule H VII) during IM. The prefrontal and parietal regions revealed no significant changes during both conditions. The results of cortical activity support the hypothesis that motor imagery and motor performance possess similar neural substrates. The differential activation in the cerebellum during EM and IM is in accordance with the assumption that the posterior cerebellum is involved in the inhibition of movement execution during imagination.
903 citations
TL;DR: It is proposed that the extent to which self-produced tactile sensation is attenuated is proportional to the error between the sensory feedback predicted by an internal forward model of the motor system and the actual sensory feedback produced by the movement.
Abstract: We investigated why self-produced tactile stimulation is perceived as less intense than the same stimulus produced externally.A tactile stimulus on the palm of the right hand was either externally produced, by a robot or self-produced by the subject. In the conditions in which the tactile stimulus was self-produced, subjects moved the arm of a robot with their left hand to produce the tactile stimulus on their right hand via a second robot. Subjects were asked to rate intensity of the tactile sensation and consistently rated self-produced tactile stimuli as less tickly, intense, and pleasant than externally produced tactile stimuli. Using this robotic setup we were able to manipulate the correspondence between the action of the subjects' left hand and the tactile stimulus on their right hand. First, we parametrically varied the delay between the movement of the left hand and the resultant movement of the tactile stimulus on the right hand. Second, we implemented varying degrees of trajectory perturbation and varied the direction of the tactile stimulus movement as a function of the direction of left-hand movement. The tickliness rating increased significantly with increasing delay and trajectory perturbation. This suggests that self-produced movements attenuate the resultant tactile sensation and that a necessary requirement of this attenuation is that the tactile stimulus and its causal motor command correspond in time and space. We propose that the extent to which self-produced tactile sensation is attenuated (i.e., its tickliness) is proportional to the error between the sensory feedback predicted by an internal forward model of the motor system and the actual sensory feedback produced by the movement.
746 citations
TL;DR: The present data corroborated the functional neuro-anatomy of word recognition systems suggested by other neuroimaging methods and described their timecourse, supporting a cascade-type process that involves different but interconnected neural modules, each responsible for a different level of processing word-related information.
Abstract: The aim of the present study was to examine the time course and scalp distribution of electrophysiological manifestations of the visual word recognition mechanism. Event-related potentials (ERPs) elicited by visually presented lists of words were recorded while subjects were involved in a series of oddball tasks. The distinction between the designated target and nontarget stimuli was manipulated to induce a different level of processing in each session (visual, phonological/phonetic, phonological/lexical, and semantic). The ERPs of main interest in this study were those elicited by nontarget stimuli. In the visual task the targets were twice as big as the nontargets. Words, pseudowords, strings of consonants, strings of alphanumeric symbols, and strings of forms elicited a sharp negative peak at 170 msec (N170); their distribution was limited to the occipito-temporal sites. For the left hemisphere electrode sites, the N170 was larger for orthographic than for nonorthographic stimuli and vice versa for the right hemisphere. The ERPs elicited by all orthographic stimuli formed a clearly distinct cluster that was different from the ERPs elicited by nonorthographic stimuli. In the phonological/phonetic decision task the targets were words and pseudowords rhyming with the French word vitrail, whereas the nontargets were words, pseudowords, and strings of consonants that did not rhyme with vitrail. The most conspicuous potential was a negative peak at 320 msec, which was similarly elicited by pronounceable stimuli but not by nonpronounceable stimuli. The N320 was bilaterally distributed over the middle temporal lobe and was significantly larger over the left than over the right hemisphere. In the phonological/lexical processing task we compared the ERPs elicited by strings of consonants (among which words were selected), pseudowords (among which words were selected), and by words (among which pseudowords were selected). The most conspicuous potential in these tasks was a negative potential peaking at 350 msec (N350) elicited by phonologically legal but not by phonologically illegal stimuli. The distribution of the N350 was similar to that of the N320, but it was broader and including temporo-parietal areas that were not activated in the "rhyme" task. Finally, in the semantic task the targets were abstract words, and the nontargets were concrete words, pseudowords, and strings of consonants. The negative potential in this task peaked at 450 msec. Unlike the lexical decision, the negative peak in this task significantly distinguished not only between phonologically legal and illegal words but also between meaningful (words) and meaningless (pseudowords) phonologically legal structures. The distribution of the N450 included the areas activated in the lexical decision task but also areas in the fronto-central regions. The present data corroborated the functional neuroanatomy of word recognition systems suggested by other neuroimaging methods and described their timecourse, supporting a cascade-type process that involves different but interconnected neural modules, each responsible for a different level of processing word-related information.
694 citations
TL;DR: This study examined the properties of the processes involved in the structural analysis of sentences using event-related brain potential measures (ERP) and showed that the early left anterior negativity was elicited and equally pronounced under both proportion conditions, suggesting that first-pass parsing processes are automatic, whereas second-pass parse processes are under participants' strategic control.
Abstract: In this study we examined the properties of the processes involved in the structural analysis of sentences using eventrelated brain potential measures (ERP). Previous research had shown two ERP components to correlate with phrase structure violations: an early left anterior negativity (ELAN), which is assumed to reflect first-pass parsing processes, and a late parietally distributed positivity (P600), assumed to reflect second-pass parsing processes. We hypothesized that the first-pass parsing processes are highly automatic, whereas second-pass parsing processes are more controlled. To test this hypothesis we varied the proportion of correct sentences and sentences containing phrase structure violations with incorrect sentences being either of a low (20p violation) or a high (80p violation) proportion. Results showed that the early left anterior negativity was elicited and equally pronounced under both proportion conditions. By contrast, the late positivity was elicited for a low proportion of incorrect sentences only.This data pattern suggests that first-pass parsing processes are automatic, whereas second-pass parsing processes are under participants' strategic control.
628 citations
TL;DR: Partially distinct cerebral circuits with the dorsal parietal pathway underlie distinct arithmetic operations in brain-lesioned patients.
Abstract: We measured cerebral activation with functional magnetic resonance imaging at 3 Tesla while eight healthy volunteers performed various number processing tasks known to be dissociable in brain-lesioned patients: naming, comparing, multiplying, or subtracting single digits. The results revealed the activation of a circuit comprising bilateral intraparietal, prefrontal, and anterior cingulate components. The extension and lateralization of this circuit was modulated by task demands. The intraparietal and prefrontal activation was more important in the right hemisphere during the comparison task and in the left hemisphere during the multiplication task and was intensely bilateral during the subtraction task. Thus, partially distinct cerebral circuits with the dorsal parietal pathway underlie distinct arithmetic operations.
580 citations
TL;DR: It is argued that the findings of ERP experiments investigating how and when the language comprehension system relates an incoming word to semantic representations of an unfolding local sentence and a wider discourse are most compatible with models of language processing in which there is no fundamental distinction between the integration of a word in its local and its global semantic context.
Abstract: In two ERP experiments we investigated how and when the language comprehension system relates an incoming word to semantic representations of an unfolding local sentence and a wider discourse. In Experiment 1, subjects were presented with short stories. The last sentence of these stories occasionally contained a critical word that, although acceptable in the local sentence context, was semantically anomalous with respect to the wider discourse (e.g., Jane told the brother that he was exceptionally slow in a discourse context where he had in fact been very quick). Relative to coherent control words (e.g., quick), these discourse-dependent semantic anomalies elicited a large N400 effect that began at about 200 to 250 msec after word onset. In Experiment 2, the same sentences were presented without their original story context. Although the words that had previously been anomalous in discourse still elicited a slightly larger average N400 than the coherent words, the resulting N400 effect was much reduced, showing that the large effect observed in stories depended on the wider discourse. In the same experiment, single sentences that contained a clear local semantic anomaly elicited a standard sentence-dependent N400 effect (e.g., Kutas & Hillyard, 1980). The N400 effects elicited in discourse and in single sentences had the same time course, overall morphology, and scalp distribution. We argue that these findings are most compatible with models of language processing in which there is no fundamental distinction between the integration of a word in its local (sentence-level) and its global (discourse-level) semantic context.
548 citations
TL;DR: Results support a state theory of hypnosis in which occipital increases in rCBF and delta activity reflect the alteration of consciousness associated with decreased arousal and possible facilitation of visual imagery.
Abstract: The neural mechanisms underlying hypnotic states and responses to hypnotic suggestions remain largely unknown and, to date, have been studied only with indirect methods. Here, the effects of hypnosis and suggestions to alter pain perception were investigated in hypnotizable subjects by using positron emission tomography (PET) measures of regional cerebral blood flow (rCBF) and electroencephalographic (EEG) measures of brain electrical activity. The experimental conditions included a restful state (Baseline) followed by hypnotic relaxation alone (Hypnosis) and by hypnotic relaxation with suggestions for altered pain unpleasantness (Hypnosis-with-Suggestion). During each scan, the left hand was immersed in neutral (35°C) or painfully hot (47°C) water in the first two conditions and in painfully hot water in the last condition. Hypnosis was accompanied by significant increases in both occipital rCBF and delta EEG activity, which were highly correlated with each other (r = 0.70, p < 0.0001). Peak increases in rCBF were also observed in the caudal part of the right anterior cingulate sulcus and bilaterally in the inferior frontal gyri. Hypnosis-related decreases in rCBF were found in the right inferior parietal lobule, the left precuneus, and the posterior cingulate gyrus. Hypnosis-with-suggestions produced additional widespread increases in rCBF in the frontal cortices predominantly on the left side. Moreover, the medial and lateral posterior parietal cortices showed suggestion-related increases overlapping partly with regions of hypnosis-related decreases. Results support a state theory of hypnosis in which occipital increases in rCBF and delta activity reflect the alteration of consciousness associated with decreased arousal and possible facilitation of visual imagery. Frontal increases in rCBF associated with suggestions for altered perception might reflect the verbal mediation of the suggestions, working memory, and topdown processes involved in the reinterpretation of the perceptual experience. These results provide a new description of the neurobiological basis of hypnosis, demonstrating specific patterns of cerebral activation associated with the hypnotic state and with the processing of hypnotic suggestions.
445 citations
TL;DR: It is argued that the current literature on prosopagnosia fails to demonstrate unequivocal evidence for a disproportionate impairment for faces as compared to nonface objects, and questions regarding neuropsychological evidence for the modularity of face recognition, as well as its theoretical and methodological foundations are raised.
Abstract: We argue that the current literature on prosopagnosia fails to demonstrate unequivocal evidence for a disproportionate impairment for faces as compared to nonface objects. Two prosopagnosic subjects were tested for the discrimination of objects from several categories (face as well as nonface) at different levels of categorization (basic, subordinate, and exemplar levels). Several dependent measures were obtained including accuracy, signal detection measures, and response times. The results from Experiments 1 to 4 demonstrate that, in simultaneous-matching tasks, response times may reveal impairments with nonface objects in subjects whose error rates only indicate a face deacit. The results from Experiments 5 and 6 show that, given limited stimulus presentation times for face and nonface objects, the same subjects may demonstrate a deacit for both stimulus categories in sensitivity. In Experiments 7, 8 and 9, a match-to-sample task that places greater demands on memory led to comparable recognition sensitivity with both face and nonface objects. Regardless of object category, the prosopagnosic subjects were more affected by manipulations of the level of categorization than normal controls. This result raises questions regarding neuropsychological evidence for the modularity of face recognition, as well as its theoretical and methodological foundations.
320 citations
TL;DR: The data suggest that the amount of activation in the various cortical regions that support visuospatial processing is related to the amount, as well as to the type, of computational demand.
Abstract: Two studies examined how the amount and type of computational demand are related to fMRI-measured activation in three bilateral cortical regions involved in the Shepard-Metzler (1971) mental-rotation paradigm. The amount of demand for the computation of visuospatial coordinates was manipulated by presenting mental rotation problems with increasing angular disparity (0, 40, 80, or 120°). Activation in both the left and right intraparietal sulcal regions increased linearly with angular disparity in two separate studies. Activation also occurred in the fusiform gyrus and inferior temporal regions, regions that are primarily associated with the processes of object and object-part identification. By contrast, the demand for object recognition and rotation processes was relatively low, and the demand for executing saccades was high in a control condition that required making a systematic visual scan of two grids. The grid-scanning condition resulted in relatively less activation in the parietal and inferior temporal regions but considerable activation in frontal areas that are associated with planning and executing saccades, including the precentral gyrus and sulcus into the posterior middle frontal region.These data suggest that the amount of activation in the various cortical regions that support visuospatial processing is related to the amount, as well as to the type, of computational demand.
TL;DR: In this article, a PET study using 15O-butanol was conducted to examine the neural correlates of reading and of the phonological conversion of legal letter strings, with or without meaning.
Abstract: Silent reading and reading aloud of German words and pseudowords were used in a PET study using (15O)butanol to examine the neural correlates of reading and of the phonological conversion of legal letter strings, with or without meaning. The results of 11 healthy, right-handed volunteers in the age range of 25 to 30 years showed activation of the lingual gyri during silent reading in comparison with viewing a fixation cross. Comparisons between the reading of words and pseudowords suggest the involvement of the middle temporal gyri in retrieving both the phonological and semantic code for words. The reading of pseudowords activates the left inferior frontal gyrus, including the ventral part of Broca's area, to a larger extent than the reading of words. This suggests that this area might be involved in the sublexical conversion of orthographic input strings into phonological output codes. (Pre)motor areas were found to be activated during both silent reading and reading aloud. On the basis of the obtained activation patterns, it is hypothesized that the articulation of high-frequency syllables requires the retrieval of their concomitant articulatory gestures from the SMA and that the articulation of low-frequency syllables recruits the left medial premotor cortex.
TL;DR: Both exogenous and endogenous orienting, but not the control condition, activated bilateral parietal and dorsal premotor regions, including the frontal eye fields, which suggest a specific role for these regions in preparatory responding to peripheral stimuli.
Abstract: Whole-brain functional magnetic resonance imaging (MRI) was used to examine the neural substrates of internally (endogenous) and externally (exogenous) induced covert shifts of attention. Thirteen normal subjects performed three orienting conditions: endogenous (location of peripheral target predicted by a central arrow 80p of the time), exogenous (peripheral target preceded by a noninformative peripheral cue), and control (peripheral target preceded by noninformative central cue). Behavioral results indicated faster reaction times (RTs) for valid than for invalid trials for the endogenous condition but slower RTs for valid than for invalid trials for the exogenous condition (inhibition of return). The spatial extent and intensity of activation was greatest for the endogenous condition, consistent with the hypothesis that endogenous orienting is more effortful (less automatic) than exogenous orienting. Overall, we did not observe distinctly separable neural systems associated with the endogenous and exogenous orienting conditions. Both exogenous and endogenous orienting, but not the control condition, activated bilateral parietal and dorsal premotor regions, including the frontal eye fields. These results suggest a specific role for these regions in preparatory responding to peripheral stimuli. The right dorsolateral prefrontal cortex (BA 46) was activated selectively by the endogenous condition. This finding suggests that voluntary, but not reflexive, shifts of attention engage working memory systems.
TL;DR: The shift in weak priming seen after REM sleep awakenings suggests that cognition during REM sleep is qualitatively different from that of waking and NREM sleep and may reflect a shift in associative memory systems, a shift that is hypothesize underlies the bizarre and hyperassociative character of REM-sleep dreaming.
Abstract: The notion that dreaming might alter the strength of associative links in memory was first proposed almost 200 years ago. But no strong evidence of such altered associative links has been obtained. Semantic priming can be used to quantify the strength of associative links between pairs of words; it is thought to measure the automatic spread of activation from a "node" representing one word to nodes representing semantically related words. Semantic priming could thus be used to test for global alterations in the strengths of associative links across the wake-sleep cycle. Awakenings from REM and nonREM (NREM) sleep produce a period of state carry-over during which performance is altered as a result of the brain's slow transition to full wakefulness, and cognitive testing in this period can provide information about the functioning of the brain during the prior sleep period. When subjects were tested across the night - before and after a night's sleep as well as immediately following forced awakenings from REM and NREM sleep-weak priming (e.g., thief-wrong) was found to be state dependent (p = 0.016), whereas strong priming (e.g., hot-cold) was not (p = 0.89). Weak primes were most effective in the presleep and REM sleep conditions and least effective in NREM and postsleep conditions. Most striking are analyses comparing weak and strong priming within each wake-sleep state. Contrary to the normal pattern of priming, subjects awakened from REM sleep showed greater priming by weak primes than by strong primes (p = 0.01). This result was seen in each of three protocols. In contrast, strong priming exceeded weak priming in NREM sleep. The shift in weak priming seen after REM sleep awakenings suggests that cognition during REM sleep is qualitatively different from that of waking and NREM sleep and may reflect a shift in associative memory systems, a shift that we hypothesize underlies the bizarre and hyperassociative character of REMsleep dreaming. Known changes in brainstem activity that control the transition into and maintenance of REM sleep provide a possible explanation of this shift.
TL;DR: The results suggest that orbito-frontal and visual regions interact in odor processing in a complementary way, depending on the task requirements.
Abstract: The functional anatomy of perceptual and semantic processings for odors was studied using positron emission tomography (PET). The first experiment was a pretest in which 71 normal subjects were asked to rate 185 odorants in terms of intensity, familiarity, hedonicity, and comestibility and to name the odorants. This pretest was necessary to select the most appropriate stimuli for the different cognitive tasks of the second experiment. The second one was a PET experiment in which 15 normal subjects were scanned using the water bolus method to measure regional cerebral blood flow (rCBF) during the performance in three conditions. In the first (perceptual) condition, subjects were asked to judge whether an odor was familiar or not. In the second (semantic) condition, subjects had to decide whether an odor corresponded to a comestible item or not. In the third (detection) condition, subjects had to judge whether the perceived stimulus was made of an odor or was just air. It was hypothetized that the three tasks were hierarchically organized from a superficial detection level to a deep semantic level. Odorants were presented with an air-flow olfactometer, which allowed the stimulations to be synchronized with breathing.Subtraction of activation images obtained between familiarity and control judgments revealed that familiarity judgments were mainly associated with the activity of the right orbito-frontal area, the subcallosal gyrus, the left inferior frontal gyrus, the left superior frontal gyrus, and the anterior cingulate (Brodmann's areas 11, 25, 47, 9, and 32, respectively). The comestibility minus familiarity comparison showed that comestibility judgments selectively activated the primary visual areas. In contrast, a decrease in rCBF was observed in these same visual areas for familiarity judgments and in the orbitofrontal area for comestibility judgments. These results suggest that orbito-frontal and visual regions interact in odor processing in a complementary way, depending on the task requirements.
TL;DR: Results indicate that retrieving previously acquired information about an object's typical color does not require reactivation of brain regions that subserve color perception.
Abstract: Positron emission tomography (PET) was used to investigate whether retrieving information about a specific object attribute requires reactivation of brain areas that mediate perception of that attribute. During separate PET scans, subjects passively viewed colored and equiluminant gray-scale Mondrians, named colored and achromatic objects, named the color of colored objects, and generated color names associated with achromatic objects. Color perception was associated with activations in the lingual and fusiform gyri of the occipital lobes, consistent with previous neuroimaging and human lesion studies.Retrieving information about object color (generating color names for achromatic objects relative to naming achromatic objects) activated the left inferior temporal,left frontal,and left posterior parietal cortices, replicating previous findings from this laboratory. When subjects generated color names for ach romatic objects relative to the low-level baseline of viewing gray-scale Mondrians, additional activations in the left fusiform/lateral occipital region were detected. However, these activations were lateral to the occipital regions associated with color perception and identical to occipital regions activated when subjects simply named achromatic objects relative to the same low-level baseline. This suggests that the occipital activations associated with retrieving color information were due to the perception of object form rather than to the top-down influence of brain areas that mediate color perception. Taken together, these results indicate that retrieving previously acquired information about an object's typical color does not require reactivation of brain regions that subserve color perception.
TL;DR: It is shown that considerable information is available from neuronal responses even under backward masking conditions that allow the neurons to have their main response in 30 msec, providing evidence for how rapid the processing of visual information is in a cortical area and provides a fundamental constraint for understanding how cortical information processing operates.
Abstract: Backward masking can potentially provide evidence of the time needed for visual processing, a fundamental constraint that must be incorporated into computational models of vision. Although backward masking has been extensively used psychophysically, there is little direct evidence for the effects of visual masking on neuronal responses. To investigate the effects of a backward masking paradigm on the responses of neurons in the temporal visual cortex, we have shown that the response of the neurons is interrupted by the mask. Under conditions when humans can just identify the stimulus, with stimulus onset asynchronies (SOA) of 20 msec, neurons in macaques respond to their best stimulus for approximately 30 msec. We now quantify the information that is available from the responses of single neurons under backward masking conditions when two to six faces were shown. We show that the information available is greatly decreased as the mask is brought closer to the stimulus. The decrease is more marked than the decrease in firing rate because it is the selective part of the firing that is especially attenuated by the mask, not the spontaneous firing, and also because the neuronal response is more variable at short SOAs. However, even at the shortest SOA of 20 msec, the information available is on average 0.1 bits. This compares to 0.3 bits with only the 16-msec target stimulus shown and a typical value for such neurons of 0.4 to 0.5 bits with a 500- msec stimulus. The results thus show that considerable information is available from neuronal responses even under backward masking conditions that allow the neurons to have their main response in 30 msec. This provides evidence for how rapid the processing of visual information is in a cortical area and provides a fundamental constraint for understanding how cortical information processing operates.
TL;DR: This experiment failed to find evidence, at the single-subject level and at the group level, for anatomical segregation of spatial and object working memory function in the frontal cortex, and revealed a sensitivity to position in the trial in the spatial, but not the object, condition.
Abstract: Behavioral studies indicate that spatial and object working memory are computed by dissociable subsystems. We investigated the neural bases of this dissociation with a whole-brain fMRI design and analysis technique that permitted direct assessment of delay-period activity, uncontaminated by other components of the trial. The task employed a ''what''-then''where'' design, with an object and a spatial delay period incorporated in each trial; within-trial order of delay conditions was balanced across each scan. Our experiment failed to find evidence, at the single-subject level and at the group level, for anatomical segregation of spatial and object working memory function in the frontal cortex. Delay-period activity in the caudate nucleus revealed a sensitivity to position in the trial in the spatial, but not the object, condition. In posterior regions, spatial delay-period activity was associated with preferential recruitment of extrastriate areas falling within Brodmann's area 19 and, less reliably,the superior parietal lobule. Object-specific delay-period activity was found predominantly in ventral regions of the posterior cortex and demonstrated more topographic variability across subjects than did spatial working memory activity.
TL;DR: This work explored the influence of type of code and mental set in two regions in the frontal cortex that have been previously shown to be involved in memory and demonstrated code-specific effects.
Abstract: The frontal cortex has been described as playing both ''setspecific'' and ''code-specific'' roles in human memory processing. Set specificity refers to the finding of goal-oriented differences in activation patterns (e.g., encoding relative to retrieval). Code specificity refers to the finding of different patterns of activation for different types of stimuli (e.g., verbal/nonverbal). Using a two (code: verbal, nonverbal) by two (set: encoding, retrieval) within-subjects design and fMRI, we explored the influence of type of code and mental set in two regions in the frontal cortex that have been previously shown to be involved in memory. A region in the dorsal extent of the inferior frontal gyrus (BA 6/44) demonstrated code-specific effects. Specifically, an interaction of material type with hemisphere was obtained, such that words produced predominantly left-lateralized activation, whereas unfamiliar faces elicited predominantly right-lateralized activation. A region of the right frontal polar cortex (in or near BA 10), which has been activated in many memory retrieval studies, showed set-specific activation in that it was more active during retrieval than encoding. These data demonstrate that distinct regions in the frontal cortex contribute in systematic yet different ways to human memory processing.
TL;DR: The evidence suggests that activation of the fusiform gyrus is most likely related to visual perceptual semantic processing, and the inferior/middle frontal lobe activity observed while performing the picture naming and semantic judgment tasks does not appear to be due to the effects of anticipation or preparation.
Abstract: A PET study of 10 normal males was carried out using the bolus H215O intravenous injection technique to examine the effects of picture naming and semantic judgment on blood flow. In a series of conditions, subjects (1) passively viewed flashing plus signs, (2) noted the occurrence of abstract patterns, (3) named animal pictures, or (4) carried out a semantic judgment on animal pictures. Anticipatory scans were carried out after the subjects were presented with the instructions but before they began the cognitive task, as they were passively viewing plus signs. Our results serve to clarify a number of current controversies regarding the neural substrate of picture naming. The results indicate that the fusiform gyrus is unlikely to be the region where low-level perceptual processing such as shape analysis is undertaken. In fact, our evidence suggests that activation of the fusiform gyrus is most likely related to visual perceptual semantic processing. In addition, the inferior/middle frontal lobe activity observed while performing the picture naming and semantic judgment tasks does not appear to be due to the effects of anticipation or preparation. Furthermore, there appears to be a set of regions (a semantic network) that becomes activated regardless of whether the subjects perform a picture naming or semantic judgment task. Finally, picture naming of animals did not activate either parietal regions or anterior inferior left temporal regions, regardless of what subtraction baseline was used.
TL;DR: Investigation of a patient, who was able to associate words and pictures on the basis of semantic relationships despite extensive damage to the left frontal, inferior parietal, and superior temporal cortices, demonstrates that activity in extrasylvian temporo-parietal and medial superior frontal regions is sufcient to perform semantic similarity judgments.
Abstract: This paper demonstrates how functional imaging studies of neuropsychological patients can provide a way of determining which areas in a cognitive network are jointly necessary and sufficient. The approach is illustrated with an investigation of the neural system underlying semantic similarity judgments. Functional neuroimaging demonstrates that normal subjects activate left temporal, parietal, and inferior frontal cortices during this task relative to physical size judgments. Neuropsychology demonstrates that damage to the temporal and parietal regions results in semantic deficits, indicating that these areas are necessary for task performance. In contrast, damage to the inferior frontal cortex does not impair task performance, indicating that the inferior frontal cortex might not be necessary. However, there are two other possible accounts of intact performance following frontal lobe damage: (1) there is functional reorganization involving the right frontal cortex and (2) there is peri-infarct activity around the damaged left-hemisphere tissue. Functional imaging of the patient is required to discount these possibilities. We investigated a patient (SW), who was able to associate words and pictures on the basis of semantic relationships despite extensive damage to the left frontal, inferior parietal, and superior temporal cortices. Although SW showed peri-infarct activation in left extrasylvian temporal cortices, no activity was observed in either left or right inferior frontal cortices. These findings demonstrate that activity in extrasylvian temporo-parietal and medial superior frontal regions is sufficient to perform semantic similarity judgments. In contrast, the left inferior frontal activations detected in each control subject appear not to be necessary for task performance. In conclusion, necessary and sufficient brain systems can be delineated by functional imaging of brain-damaged patients who are not functionally impaired.
TL;DR: This work evaluated the first direct neural support for mapping the subitizing-counting dichotomy onto separable processes mediating preattentive vision and shifts of visual attention at a neural level.
Abstract: Visual object enumeration is rapid and accurate for four or fewer items but slow and error-prone for over four items. This dichotomy has recently been linked to visual attentional phenomena by findings suggesting that ''subitizing'' of small sets of objects is preattentive whereas ''counting'' of over four items demands spatial shifts of attention. We evaluated this link at a neural level, using H215O positron emission tomography to measure changes in regional cerebral blood flow while subjects enumerated the number of target vertical bars that ''popped out'' of a 16-bar visual display consisting of both horizontal and vertical bars. Relative to a condition with a single target, subi tizing (one to four targets) activated foci in the occipital extrastriate cortex, consistent with involvement of early,preattentive visual processes. Relative to subitizing, counting (five to eight targets) activated a widespread network of brain regions, including multiple foci implicated in shifting visual attention- large regions of the superior parietal cortex bilaterally and a focus in the right inferior frontal cortex.These results offer the first direct neural support for mapping the subitizing-counting dichotomy onto separable processes mediating preattentive vision and shifts of visual attention.
TL;DR: It is suggested that the brain honors the distinction between open-and closed-class words, in relation to the different roles that they play in on-line sentence processing.
Abstract: This paper presents evidence of the disputed existence of an electrophysiological marker for the lexical-categorical distinction between open- and closed-class words. Event-related brain potentials were recorded from the scalp while subjects read a story. Separate waveforms were computed for open- and closed-class words. Two aspects of the waveforms could be reliably related to vocabulary class. The first was an early negativity in the 230- to 350-msec epoch, with a bilateral anterior predominance. This negativity was elicited by open- and closedclass words alike, was not affected by word frequency or word length, and had an earlier peak latency for closed-class words. The second was a frontal slow negative shift in the 350- to 500-msec epoch, largest over the left side of the scalp. This late negativity was only elicited by closed-class words. Although the early negativity cannot serve as a qualitative marker of the open- and closed-class distinction, it does reflect the earliest electrophysiological manifestation of the availability of categorical information from the mental lexicon. These results suggest that the brain honors the distinction between open- and closed-class words, in relation to the different roles that they play in on-line sentence processing.
TL;DR: Whereas control subjects perceived words as wholes, dyslexic subjects may have relied on sublexical word recognition and occasionally mistook a correctly beginning word for the one they had expected, as well as to both orthographically and semantically inappropriate sentence-ending words.
Abstract: The combined temporal and spatial resolution of MEG (magnetoencephalography) was used to study whether the same brain areas are similarly engaged in reading comprehension in normal and developmentally dyslexic adults. To extract a semantically sensitive stage of brain activation we manipulated the appropriateness of sentence-ending words to the preceding sentence context. Sentences, presented visually one word at a time,either ended with a word that was (1) expected, (2) semantically appropriate but unexpected, (3) semantically anomalous but sharing the initial letters with the expected word, or (4) both semantically and orthographically inappropriate to the sentence context. In both subject groups all but the highly expected sentence endings evoked strong cortical responses, localized most consistently in the left superior temporal cortex, although additional sources were occasionally found in more posterior parietal and temporal areas and in the right hemisphere. Thus, no significant differences were found in the spatial distribution of brain areas involved in semantic processing between fluent and dyslexic readers. However,both timing and strength of activation clearly differed between the two groups.First, activation sensitivity to word meaning within a sentence context began about 100 msec later in dyslexic than in control subjects. This is likely to result from affected presemantic processing stages in dyslexic readers. Second, the neural responses were significantly weaker in dyslexic than in control subjects, indicating involvement of a smaller or lesssynchronous neural population in reading comprehension. Third, in contrast to control subjects, the dyslexic readers showed significantly weaker activation to semantically inappropriate words that began with the same letters as the most expected word than to both orthographically and semantically inappropriate sentence-ending words. Thus, word recognition by the dyslexic group seemed to be qualitatively different: Whereas control subjects perceived words as wholes, dyslexic subjects may have relied on sublexical word recognition and occasionally mistook a correctly beginning word for the one they had expected.
TL;DR: The results suggest that the visual processing capacity of deaf children and adolescents does not exceed that of age- and gender-matched hearing subjects, but deaf school children show deficits in visual processing in conditions with and without attentional load.
Abstract: The research concerning the visual perception in deaf subjects has led to contradictory results: Deaf subjects have been reported to show enhanced visual perceptual skills compared to hearing subjects (Neville & Lawson, 1987). On the other hand, there are indications that acoustic deprivation may produce an inferiority in all sensory modalities (Myklebust, 1964). These contradictions may be due to methodological differences: The investigators selected different conditions (e.g. attentive/nonattentive) and various samples of deaf subjects (e.g., different age, language, and aetiology groups). In our study, we tested a large sample of deaf subjects with texture segmentation and visual search conditions, which allowed us to differentiate between visual processing with and without attentional load. All deaf subjects had profound hearing loss within the first year of life. Our results suggest that the visual processing capacity of deaf children and adolescents does not exceed that of age- and gender-matched hearing subjects. Rather, deaf school children show deficits in visual processing in conditions with and without attentional load. Age (6 to 20 years), language used (oral, sign, oral + sign), and aetiology for deafness (genetic, maternal rubella, perinatal, infection in the first year of life, unknown) did not consistently influence the results. The deficits in visual processing were partially compensated for in adult deaf subjects. The performances of deaf and hearing adults in trials that could be solved preattentively did not differ statistically significantly, but in attention-dependent trials the deaf subjects were more efficient than the hearing controls. We conclude that visual compensation for deafness is limited to attention-dependent tasks and does not develop until adulthood.
TL;DR: Controls and nontactile-extinction groups performed better when the hands were in anatomical than in crossed position, and patients with tactile extinction detected contralesional stimuli with higher accuracy in crossed than in anatomical position suggests that impairments in detecting contralsional stimuli can be due not only to sensory but also to spatial factors contingent upon the position of the hands.
Abstract: Twelve normal controls, twelve left-brain-damaged patients, and thirty-six right-brain-damaged patients with or without tactile extinction or tactile neglect were asked to report light touches delivered to the left or the right hand or simultaneously to both hands. The hands could be in anatomic position or one hand could cross over the other. Moreover, the two hands could be in the left or the right hemispace or across the corporeal midline. Controls and nontactile-extinction groups performed better when the hands were in anatomical than in crossed position. By contrast, patients with tactile extinction detected contralesional stimuli with higher accuracy in crossed than in anatomical position. This result suggests that, in these patients, impairments in detecting contralesional stimuli can be due not only to sensory but also to spatial factors contingent upon the position of the hands. There was no interaction between the effect of crossing the hands and the hemispace where the crossing took place. This suggests that coding the position of a hand as left or right does not necessarily occur in relation to the bodily midline, but it may arise from the computation of the position of the other hand.
TL;DR: Recorded intracranial ERPs from various structures during a continuous recognition memory task provide electrophysiological evidence of multiple anatomo-functional memory systems in the human brain: a short-term semantic activation system and a long-term episodic memory system, with interface structures that coordinate the functioning of these two systems.
Abstract: Prior exposure to a stimulus can facilitate the performance to subsequent presentations of that stimulus. ERP studies have shown that this facilitation is associated with the modulation of two components (N400 and P600). Investigation of the time course of both behavioral and ERP repetition effects have led to the assumption that it is subserved by the combination of at least two mechanisms operating at different time-points: a short-delay semantic activation and a long-lasting episodic mechanism. The present experiment recorded intracranial ERPs from various structures during a continuous recognition memory task to investigate the respective contribution of the different brain regions to short- and long-delay ERP repetition effects. The results are in good agreement with both the classical neuropsychological literature and the more recent data obtained with functional imagery techniques. They provide electrophysiological evidence of multiple anatomo-functional memory systems in the human brain: a short-term semantic activation system and a long-term episodic memory system, with interface structures that coordinate the functioning of these two systems.
TL;DR: A novel dependent measure was introduced to event-related potential research the latency of spontaneous, posttrial blinking, which suggests that an earlier phase of response selection is influenced by accessory stimulation.
Abstract: When an intense but task-irrelevant "accessory" stimulus accompanies the imperative stimulus in choice reaction task, reaction times (RTs) are facilitated. In a similar previous study (Hackley & Valle-Inclan,1998), we showed that this effect is not due to a reduction of the interval from onset of the lateralized readiness potential (LRP) until movement onset. In the present study, the RT task was modified to move a portion of the response selection stage into this time interval. The interval remained invariant, indicating that this late phase of the response selection process is not speeded by a accessory stimulation. However, we observed amplitude modulation of the LRP on no-go trials in a condition with three alternative responses. This finding suggests that an earlier phase of response selection is influenced by accessory stimulation. In addition, a novel dependent measure was introduced to event-related potential research-the latency of spontaneous, posttrial blinking.
TL;DR: It is shown that reduced activity in the extrastriate cortex accompanies within-modality priming in both visual and auditory modalities, and this work suggests that priming-related decreases in this region are associated with more general aspects of priming.
Abstract: Previous neuroimaging studies of perceptual priming have reported priming-related decreases in the extrastriate cortex. However, because these experiments have used visual stimuli, it is unclear whether the observed decreases are associated specifically with some aspect of visual perceptual processing or with more general aspects of priming. We studied within-and cross-modality priming using an auditory word stem completion paradigm. Positron emission tomography (PET) images were obtained during stem completion and a fixation task. Within-modality auditory priming was associated with blood flow decreases in the extrastriate cortex (bilateral), medial/right anterior prefrontal cortex, right angular gyrus, and precuneus. In cross-modality priming, the study list was presented visually, and subjects completed auditory word stems. Cross-modality priming was associated with trends for blood flow decreases in the left angular gyrus and increases in the medial/right anterior prefrontal cortex. Results thus indicate that reduced activity in the extrastriate cortex accompanies within-modality priming in both visual and auditory modalities.
TL;DR: The data suggest that the attentional demands of this task are relatively greater for older adults and consequently lead to the recruitment of additional neural systems during task performance.
Abstract: We used H215O positron emission tomography (PET) to measure age-related changes in regional cerebral blood flow (rCBF) during a verbal recognition memory task. Twelve young adults (20 to 29 years) and 12 older adults (62 to 79 years) participated. Separate PET scans were conducted during Encoding, Baseline, and Retrieval conditions. Each of the conditions involved viewing a series of 64 words and making a two-choice response manually. The complete reaction time (RT) distributions in each task condition were characterized in terms of an ex-Gaussian model (convolution of exponential and Gaussian functions). Parameter estimates were obtained for the mean of the exponential component (τ), representing a task-specific decision process and the mean of the Gaussian component (μ), representing residual sensory coding and response processes. Independently of age group, both μ and τ were higher in the Encoding and Retrieval conditions than in the Baseline condition, and τ was higher during Retrieval than during Encoding. Age-related slowing in task performance was evident primarily in μ. For young adults, rCBF activation in the right prefrontal cortex, in the Retrieval condition, was correlated positively with μ but not with τ. For older adults, rCBF changes (both increases and decreases) in several cortical regions were correlated with both μ and τ. The data suggest that the attentional demands of this task are relatively greater for older adults and consequently lead to the recruitment of additional neural systems during task performance.