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Language-related field potentials in the anterior-medial temporal lobe: I. Intracranial distribution and neural generators.
Field potentials were recorded from intracranial electrodes in humans to study language-related processing. Subjects viewed sentences in which each word was presented successively in the center of a video monitor. Half of the sentences ended normally, while the other half ended with a semantically anomalous word. The anomalous sentence-ending words elicited a large negative field potential with a peak latency near 400 msec, which was focally distributed bilaterally in the anterior medial temporal lobe (AMTL), anterior to the hippocampus and near the amygdala. Subdural electrodes positioned near the collateral sulcus just inferior and lateral to the amygdala recorded a positive field potential at the same latency. This spatial distribution of voltage suggested that this language-sensitive field potential was generated in the neocortex near the collateral sulcus and anterior fusiform gyrus. Additional task-related field potentials were recorded in the hippocampus. The AMTL field potential at 400 msec shares characteristics with the N400 potential recorded from scalp electrodes that has been associated with semantic processing.
Language-related field potentials in the anterior-medial temporal lobe: II. Effects of word type and semantic priming.
Field potentials were recorded from intracranial electrodes in humans to study the role of the anterior medial temporal lobe (AMTL) in language-related processing. Subjects viewed lists of words in which orthography and word type varied, or in which words were primed by semantic associates. Large negative field potentials were elicited within the AMTL by isolated words. The amplitude and intracranial distribution of these AMTL field potentials were consistent with those in our previous study in which anomalous sentence-ending words were used as stimuli (McCarthy et al., in press). The neocortex, in the region of the collateral sulcus and anterior fusiform gyrus, was identified as the likely neural generator of this field potential. The AMTL field potential was diminished by semantic priming, and was larger for words with semantic content than for words serving grammatical function. Orthographically illegal nonwords did not elicit this field potential. The N400 scalp event-related potential (ERP) has been shown to respond in the same manner to these task manipulations (Nobre and McCarthy, 1994), and, thus, the AMTL field potential was proposed to contribute to the generation of N400. The possible roles in language processing reflected by the AMTL field potential were considered.
Face recognition in human extrastriate cortex.
1. Twenty-four patients with electrodes chronically implanted on the surface of extrastriate visual cortex viewed faces, equiluminant scrambled faces, cars, scrambled cars, and butterflies. 2. A surface-negative potential, N200, was evoked by faces but not by the other categories of stimuli. N200 was recorded only from small regions of the left and right fusiform and inferior temporal gyri. Electrical stimulation of the same region frequently produced a temporary inability to name familiar faces. 3. The results suggest that discrete regions of inferior extrastriate visual cortex, varying in location between individuals, are specialized for the recognition of faces. These "face modules" appear to be intercalated among other functionally specific small regions.
Orienting attention to locations in mental representations.
Many cognitive processes depend on our ability to hold information in mind, often well beyond the offset of the original sensory input. The capacity of this visual short-term memory (VSTM) is limited to around three to four items. Recent research has demonstrated that the content of VSTM can be modulated by top-down attentional biases. This has been demonstrated using retrodictive spatial cues, termed "retro-cues," which orient subjects' attention to spatial locations within VSTM. In the present article, we tested whether the use of these cues is modulated by memory load and cue delay. There are a number of important conclusions: (1) Top-down biases can operate on very brief iconic traces as well as on older VSTM representations (Exp. 1). (2) When operating within capacity, subjects use the cue to prioritise where they initiate their memory search, rather than to discard uncued items (Exps. 2 and 3). (3) When capacity is exceeded, there is little benefit to top-down biasing relative to a neutral condition; however, unattended items are lost, with there being a substantial cost of invalid spatial cueing (Exp. 3). (4) These costs and benefits of orienting spatial attention differ across iconic memory and VSTM representations when VSTM capacity is exceeded (Exp. 4).
Brain activations during visual search: contributions of search efficiency versus feature binding.
We investigated the involvement of the parietal cortex in binding features during visual search using functional magnetic resonance imaging. We tested 10 subjects in four visual search tasks across which we independently manipulated (1) the requirement to integrate different types of features in a stimulus (feature or conjunction search) and (2) the degree of search efficiency (efficient or inefficient). We identified brain areas that were common to all conditions of visual search and areas that were sensitive to the factors of efficiency and feature binding. Visual search engaged an extensive network of parietal, frontal, and occipital areas. The factor of efficiency exerted a strong influence on parietal activations along the intraparietal sulcus and in the superior parietal lobule. These regions showed a main effect of efficiency and showed a simple effect when inefficient conditions were compared directly with efficient pop-out conditions in the absence of feature binding. Furthermore, a correlation analysis supported a tight correspondence between posterior parietal activation and the slope of reaction-time search functions. Conversely, feature binding during efficient pop-out search was not sufficient to modulate the parietal cortex. The results confirm the important role of the parietal cortex in visual search, but suggest that feature binding is not a requirement to engage its contribution.
The noradrenergic alpha2 agonist clonidine modulates behavioural and neuroanatomical correlates of human attentional orienting and alerting.
We examined whether the known noradrenergic attenuation of the alerting effect (the beneficial effect of a warning cue) results from an underlying effect of noradrenaline on temporal orienting (orienting toward a particular moment in time). Following a within-subjects, counterbalanced design, 10 healthy human volunteers received placebo, 200 microg clonidine or 1 mg guanfacine (alpha2 agonists) in three separate testing sessions. Subjects were scanned by fMRI while performing attentional orienting tasks containing spatially informative, temporally informative, non-informative or no cues. The alerting effect primarily activated left-lateralized prefrontal, premotor and parietal regions. Clonidine, but not guanfacine, impaired behavioural measures of the alerting effect while attenuating activity in the left temporo-parietal junction. Replicating previous results, the temporal orienting task activated left parietal and frontal cortex, while parietal cortex was activated bilaterally during spatial orienting. Of these networks, clonidine, but not guanfacine, attenuated left prefrontal cortex and insula activity during temporal orienting and attenuated right superior parietal cortex activity during spatial orienting,. To complement these neuroanatomical changes, clonidine produced selective behavioural effects on both temporal and spatial orienting. The anatomical dissociation between the effects of clonidine during temporal orienting versus alerting suggests that noradrenergic modulation of the alerting effect does not result only from an underlying effect on temporal orienting. Furthermore, we have demonstrated lateralized neuroanatomical substrates for the noradrenergic modulation of human attentional orienting in the spatial and temporal domains.
Dissociating linguistic processes in the left inferior frontal cortex with transcranial magnetic stimulation.
Is the left inferior frontal cortex (LIFC) a single functional region, or can it be subdivided into distinct areas that contribute differently to word processing? Here we used transcranial magnetic stimulation (TMS) to investigate anterior and posterior LIFC when the meaning and sound of words were being processed. Relative to no stimulation, TMS of the anterior LIFC selectively increased response latencies when participants focused on the meaning of simultaneously presented words (i.e., synonym judgments) but not when they focused on the sound pattern of the words (i.e., homophone judgments). In contrast, the opposite dissociation was observed in the posterior LIFC, where stimulation selectively interfered with the phonological but not the semantic task. This double dissociation shows functionally distinct subdivisions of the LIFC that can be understood in terms of separable corticocortical connections linking the anterior LIFC to temporal pole regions associated with semantic memory and the posterior LIFC to temporoparietal regions involved in auditory speech processing.
Functional magnetic resonance imaging of human prefrontal cortex activation during a spatial working memory task.
High-speed magnetic resonance (MR) imaging was used to detect activation in the human prefrontal cortex induced by a spatial working memory task modeled on those used to elucidate neuronal circuits in nonhuman primates. Subjects were required to judge whether the location occupied by the current stimulus had been occupied previously over a sequence of 14 or 15 stimuli presented in various locations. Control tasks were similar in all essential respects, except that the subject's task was to detect when one of the stimuli presented was colored red (color detection) or when a dot briefly appeared within the stimulus (dot detection). In all tasks, two to three target events occurred randomly. The MR signal increased in an area of the middle frontal gyrus corresponding to Brodmann's area 46 in all eight subjects performing the spatial working memory task. Right hemisphere activation was greater and more consistent than left. The MR signal change occurred within 6-9 sec of task onset and declined within a similar period after task completion. An increase in MR signal was also noted in the control tasks, but the magnitude of change was less than that recorded in the working memory task. These differences were replicated when testing was repeated in five of the original subjects. The localization of spatial working memory function in humans to a circumscribed area of the middle frontal gyrus supports the compartmentalization of working memory functions in the human prefrontal cortex and the localization of spatial memory processes to comparable areas in humans and nonhuman primates.
Anticipating conflict facilitates controlled stimulus-response selection.
Cognitive control can be triggered in reaction to previous conflict, as suggested by the finding of sequential effects in conflict tasks. Can control also be triggered proactively by presenting cues predicting conflict ("proactive control")? We exploited the high temporal resolution of ERPs and controlled for sequential effects to ask whether proactive control based on anticipating conflict modulates neural activity related to cognitive control, as may be predicted from the conflict-monitoring model. ERPs associated with conflict detection (N2) were measured during a cued flanker task. Symbolic cues were either informative or neutral with respect to whether the target involved conflicting or congruent responses. Sequential effects were controlled by analyzing the congruency of the previous trial. The results showed that cueing conflict facilitated conflict resolution and reduced the N2 latency. Other potentials (frontal N1 and P3) were also modulated by cueing conflict. Cueing effects were most evident after congruent than after incongruent trials. This interaction between cueing and sequential effects suggests neural overlap between the control networks triggered by proactive and reactive signals. This finding clarifies why previous neuroimaging studies, in which reactive sequential effects were not controlled, have rarely found anticipatory effects upon conflict-related activity. Finally, the high temporal resolution of ERPs was critical to reveal a temporal modulation of conflict detection by proactive control. This novel finding suggests that anticipating conflict speeds up conflict detection and resolution. Recent research suggests that this anticipatory mechanism may be mediated by preactivation of ACC during the preparatory interval.
Searching for targets within the spatial layout of visual short-term memory.
Recent studies have revealed that the internal representations that we construct from the environment and maintain in visual short-term memory (VSTM) to guide behavior are highly flexible and can be selectively modulated according to our task goals and expectations. In the current study, we conducted two experiments to compare and contrast neural mechanisms of selective attention related to searching for target items within perceptual versus VSTM representations. We used event-related potentials to investigate whether searching for relevant target items from within VSTM representations involves spatially specific biasing of neural activity in a manner analogous to that which occurs during visual search for target items in perceptual arrays. The results, replicated across the two experiments, revealed that selection of a target object within a search array maintained in VSTM proceeds through a similar mechanism as that in the perceptual domain. In line with previous results, N2pc potentials were obtained when targets were identified within a perceptual visual-search array. Interestingly, equivalent N2pcs, with similar time courses and scalp distributions, were also elicited when target items were identified within a VSTM representation. The findings reinforce the notion of highly flexible VSTM representations that can be modulated according to task goals and suggest a large degree of overlap in the spatially specific neural mechanisms of target selection across the perceptual and VSTM domains.
Auditory evoked visual awareness following sudden ocular blindness: an EEG and TMS investigation.
Neurophysiological and neuroanatomical studies have provoked controversy about whether the visual cortex may be more modifiable than previously believed. Auditory processing is enhanced in blind compared to sighted people, and the enhancement might reflect encroachment of auditory transmission onto visual cortex. To address this issue, we recorded the auditory event-related potentials (ERPs) correlated with auditory related paradoxical visual awareness in a subject with traumatic total late-onset blindness. We found that (1) there was auditory related brain activity over the occipital visual scalp regions starting from a very early stage (< 80 ms) and (2) this occipital activity was significantly different between "visually aware" and "visually unaware" responses in the P1 (40-80 ms) component following meaningful stimuli. There was also a significant difference between responses with and without visual awareness in the N1 (100-120 ms) component following either tones or meaningful stimuli. The phosphenes accompanying auditory stimuli in the ERP experiment were always perceived to be directly in front of the subject and this was reproduced by transcranial magnetic stimulation over the blind primary visual cortex and by sudden sounds delivered to the side or behind the subject. The TMS induced phosphenes were restricted to the central part of the space and were, at least qualitatively, the same as those induced by sounds. The results are clear evidence that human perceptual functions can be reorganized after sudden, late-onset, total ocular blindness.
Biasing perception by spatial long-term memory.
Human perception is highly flexible and adaptive. Selective processing is tuned dynamically according to current task goals and expectations to optimize behavior. Arguably, the major source of our expectations about events yet to unfold is our past experience; however, the ability of long-term memories to bias early perceptual analysis has remained untested. We used a noninvasive method with high temporal resolution to record neural activity while human participants detected visual targets that appeared at remembered versus novel locations within naturalistic visual scenes. Upon viewing a familiar scene, spatial memories changed oscillatory brain activity in anticipation of the target location. Memory also enhanced neural activity during early stages of visual analysis of the target and improved behavioral performance. Both measures correlated with subsequent target-detection performance. We therefore demonstrated that memory can directly enhance perceptual functions in the human brain.
Word recognition in the human inferior temporal lobe.
Studies of primates and of patients with brain lesions have shown that the visual system represents the external world in regions and pathways specialized to compute visual features and attributes. For example, object recognition is performed by a ventral pathway located in the inferior portion of the temporal lobe. We studied visual processing of words and word-like stimuli (letter-strings) by recording field potentials directly from the human inferior temporal lobe. Our results showed that two discrete portions of the fusiform gyrus responded preferentially to letter-strings. A region of the posterior fusiform gyrus responded equally to words and non-words, and was unaffected by the semantic context in which words were presented. In contrast, a region of the anterior fusiform gyrus was sensitive to these stimulus dimensions. These regions were distinct from areas that responded to other types of complex visual stimuli, including faces and coloured patterns, and thus form a functionally specialized stream within the ventral visual pathway.
Task-switching and memory retrieval processing: electrophysiological evidence.
Participants attempted to retrieve phonological or imagery-based information under conditions where either the two kinds of retrieval were required in separate blocks, or where frequent switches between retrieval tasks were required within blocks. Electrophysiological indices of processes engaged in pursuit of accurate memory judgments, elicited by contrasting ERPs evoked by correctly identified new test items, differed according to retrieval task only when the tasks were completed in separate blocks. The principal conclusion suggested by these findings is that the requirement to alternate frequently between the two retrieval tasks attenuated the engagement of task-specific processes that form part of a retrieval attempt.
Orienting attention based on long-term memory experience.
Attentional orienting and memory are intrinsically bound, but their interaction has rarely been investigated. Here we introduce an experimental paradigm using naturalistic scenes to investigate how long-term memory can guide spatial attention and thereby enhance identification of events in the perceptual domain. In the task, stable memories of objects embedded within complex scenes guide spatial orienting. We compared the behavioral effects and neural systems of memory-guided orienting with those in a more traditional attention-orienting task in which transient spatial cues guide attention. Memory-guided attention operated within surprisingly short intervals and conferred reliable and sizeable advantages for detection of objects embedded in scenes. Event-related functional magnetic resonance imaging showed that memory-guided attention involves the interaction between brain areas participating in retrieval of memories for spatial context with the parietal-frontal network for visual spatial orienting. Activity in the hippocampus was specifically engaged in memory-guided spatial attention and correlated with the ensuing behavioral advantage.
Orienting attention to locations in internal representations.
Three experiments investigated whether it is possible to orient selective spatial attention to internal representations held in working memory in a similar fashion to orienting to perceptual stimuli. In the first experiment, subjects were either cued to orient to a spatial location before a stimulus array was presented (pre-cue), cued to orient to a spatial location in working memory after the array was presented (retro-cue), or given no cueing information (neutral cue). The stimulus array consisted of four differently colored crosses, one in each quadrant. At the end of a trial, a colored cross (probe) was presented centrally, and subjects responded according to whether it had occurred in the array. There were equivalent patterns of behavioral costs and benefits of cueing for both pre-cues and retro-cues. A follow-up experiment used a peripheral probe stimulus requiring a decision about whether its color matched that of the item presented at the same location in the array. Replication of the behavioral costs and benefits of pre-cues and retro-cues in this experiment ruled out changes in response criteria as the only explanation for the effects. The third experiment used event-related potentials (ERPs) to compare the neural processes involved in orienting attention to a spatial location in an external versus an internal spatial representation. In this task, subjects responded according to whether a central probe stimulus occurred at the cued location in the array. There were both similarities and differences between ERPs to spatial cues toward a perception versus an internal spatial representation. Lateralized early posterior and later frontal negativities were observed for both pre- and retro-cues. Retro-cues also showed additional neural processes to be involved in orienting to an internal representation, including early effects over frontal electrodes.
Components of switching intentional set.
Despite the intuition that we can shift cognitive set on instruction, some behavioral studies have suggested that set shifting might only be accomplished once we engage in performance of the new task. It is possible that set switching consists of more than one component cognitive process and that the component processes might segregated in time. We recorded event-related potentials (ERPs) during two set-switching tasks to test whether different component processes were responsible for (i) set initiation and reconfiguration when presented with the instruction to switch, and (ii) the implementation of the new set once subjects engaged in performing the new task. The response switching (RS) task required shifts of intentional set; subjects selected between responses according to one of two conflicting intentional sets. The results demonstrated the existence of more than one constituent process. Some of the processes were linked to the initiation and reconfiguration of the set prior to actual performance of the new task. Other processes were time locked to performance of new task items. Set initiation started with modulation of medial frontal ERPs and was followed by modulation over parietal electrodes. Implementation of intentional set was associated with modulation of response-related ERPs.
Where and when to pay attention: the neural systems for directing attention to spatial locations and to time intervals as revealed by both PET and fMRI.
Although attention is distributed across time as well as space, the temporal allocation of attention has been less well researched than its spatial counterpart. A temporal analog of the covert spatial orientation task [Posner MI, Snyder CRR, Davidson BJ (1980) Attention and the detection of signals. J Exp Psychol Gen 109:160-174] was developed to compare the neural systems involved in directing attention to spatial locations versus time intervals. We asked whether there exists a general system for allocating attentional resources, independent of stimulus dimension, or whether functionally specialized brain regions are recruited for directing attention toward spatial versus temporal aspects of the environment. We measured brain activity in seven healthy volunteers by using positron emission tomography (PET) and in eight healthy volunteers by using functional magnetic resonance imaging (fMRI). The task manipulated cued attention to spatial locations (S) and temporal intervals (T) in a factorial design. Symbolic central cues oriented subjects toward S only (left or right), toward T only (300 msec or 1500 msec), toward both S and T simultaneously, or provided no information regarding S or T. Subjects also were scanned during a resting baseline condition. Behavioral data showed benefits and costs for performance during temporal attention similar to those established for spatial attention. Brain-imaging data revealed a partial overlap between neural systems involved in the performance of spatial versus temporal orientation of attention tasks. Additionally, hemispheric asymmetries revealed preferential right and left parietal activation for spatial and temporal attention, respectively. Parietal cortex was activated bilaterally by attending to both dimensions simultaneously. This is the first direct comparison of the neural correlates of attending to spatial versus temporal cues.
Language network specializations: an analysis with parallel task designs and functional magnetic resonance imaging.
Although the classical core regions of the language system (Broca's and Wernicke's areas) were defined over a century ago, it took the advent of functional imaging to sharpen our understanding of how these regions and adjacent parts of the brain are associated with particular aspects of language. One limitation of such studies has been the need to compare results across different subject groups, each performing a different type of language task. Thus, this study was designed to examine overlapping versus segregated brain activations associated with three fundamental language tasks, orthography, phonology and semantics performed by the same subjects during a single experimental session. The results demonstrate a set of primarily left-sided core language regions in ventrolateral frontal, supplementary motor, posterior mid-temporal, occipito-temporal and inferior parietal areas, which were activated for all language tasks. Segregated task-specific activations were demonstrated within the ventrolateral frontal, mid-temporal and inferior parietal areas. Within the inferior frontal cortex (Broca's regional complex), segregated activations were seen for the semantic and phonological tasks. These findings demonstrate both common and task specific activations within the language system.