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  • FEF TMS affects visual cortical activity.

    24 October 2018

    We tested whether the frontal eye field (FEF) is critical in controlling visual processing in posterior visual brain areas during the orienting of spatial attention. Short trains (5 pulses at 10 Hz) of transcranial magnetic stimulation (TMS) were applied to the right FEF during the cueing period of a covert attentional task while event-related potentials (ERPs) were simultaneously recorded from lateral posterior electrodes, where visual components are prominent. FEF TMS significantly affected the neural activity evoked by visual stimuli, as well as the ongoing neural activity recorded during earlier anticipation of the visual stimuli. The effects of FEF TMS started earlier and were greatest for brain activity recorded ipsilaterally to FEF TMS and contralaterally to the visual stimulus. The TMS-induced effect on visual ERPs occurred at the same time as ERPs were shown to be modulated by visual attention. Importantly, no similar effects were observed after TMS of a control site that was physically closer but not anatomically interconnected to the recording sites. The results show that the human FEF has a causal influence over the modulation of visual activity in posterior areas when attention is being allocated.

  • Cortical activation in the human brain during lateral saccades using EPISTAR functional magnetic resonance imaging.

    24 October 2018

    The location of the human cortical substrate underlying simple horizontal saccadic eye movements was investigated using echoplanar functional magnetic resonance imaging (fMRI) in young healthy volunteers. Echoplanar imaging with signal targeting and alternating radiofrequency (EPISTAR), a novel perfusion technique, measured signal intensity changes in one to four contiguous 10-mm slices centered to include both striate cortex and putative frontal eye fields during horizontal saccade and fixation conditions. Subtraction images of self-paced visually guided saccadic versus fixation conditions showed bilateral marked and statistically significant localized signal increases in the precentral region (Brodmann areas 4, 6) and peristriate cortex (areas 17, 18, 19) and qualitative increases in the superior medial frontal region, as identified by a Talairach-Tournoux generalized template in the brain slices that were scanned. Additional parietal activation occurred during a target-guided saccade task. Our data (i) support the localization of the human FEF, as identified by simple, nonexploratory saccadic eye movements, in the precentral motor strip and premotor cortex, (ii) show individual variability in the exact anatomical location of saccade-related activations, and (iii) confirm that the EPISTAR technique can demonstrate localized signal increases during a behavioral task.

  • Language-related field potentials in the anterior-medial temporal lobe: I. Intracranial distribution and neural generators.

    24 October 2018

    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.

    24 October 2018

    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.

    24 October 2018

    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.

    24 October 2018

    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).

  • Temporal order is coded temporally in the brain: early event-related potential latency shifts underlying prior entry in a cross-modal temporal order judgment task.

    24 October 2018

    The speeding-up of neural processing associated with attended events (i.e., the prior-entry effect) has long been proposed as a viable mechanism by which attention can prioritize our perception and action. In the brain, this has been thought to be regulated through a sensory gating mechanism, increasing the amplitudes of early evoked potentials while leaving their latencies unaffected. However, the majority of previous research has emphasized speeded responding and has failed to emphasize fine temporal discrimination, thereby potentially lacking the sensitivity to reveal putative modulations in the timing of neural processing. In the present study, we used a cross-modal temporal order judgment task while shifting attention between the visual and tactile modalities to investigate the mechanisms underlying selective attention electrophysiologically. Our results indicate that attention can indeed speed up neural processes during visual perception, thereby providing the first electrophysiological support for the existence of prior entry.

  • Brain activations during visual search: contributions of search efficiency versus feature binding.

    24 October 2018

    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.

    24 October 2018

    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.