Anticipatory biasing of visuospatial attention indexed by retinotopically specific alpha-band electroencephalography increases over occipital cortex.

Alpha-band (8-14 Hz) oscillatory EEG activity was examined with high-density scalp electrical recording during the cue-stimulus interval of an endogenous spatial cueing paradigm. In different blocks, cued spatial locations (left or right) were in either the upper or lower visual field, and attended stimuli were either oriented Ts or moving dots. Distractor stimuli were equally likely in the uncued hemifield. Sustained focal increases of alpha-band activity were seen over occipital cortex contralateral to the direction of the to-be-ignored location (ipsilateral to the cued direction of attention) before onset of the to-be-attended stimulus. The focus of alpha-band activity also moved depending on whether cued locations were in the upper or lower field. Results are consistent with active gating of uncued spatial locations.

Spatiotemporal activity of a cortical network for processing visual motion revealed by MEG and fMRI.

A sudden change in the direction of motion is a particularly salient and relevant feature of visual information. Extensive research has identified cortical areas responsive to visual motion and characterized their sensitivity to different features of motion, such as directional specificity. However, relatively little is known about responses to sudden changes in direction. Electrophysiological data from animals and functional imaging data from humans suggest a number of brain areas responsive to motion, presumably working as a network. Temporal patterns of activity allow the same network to process information in different ways. The present study in humans sought to determine which motion-sensitive areas are involved in processing changes in the direction of motion and to characterize the temporal patterns of processing within this network of brain regions. To accomplish this, we used both magnetoencephalography (MEG) and functional magnetic resonance imaging (fMRI). The fMRI data were used as supplementary information in the localization of MEG sources. The change in the direction of visual motion was found to activate a number of areas, each displaying a different temporal behavior. The fMRI revealed motion-related activity in areas MT+ (the human homologue of monkey middle temporal area and possibly also other motion sensitive areas next to MT), a region near the posterior end of the superior temporal sulcus (pSTS), V3A, and V1/V2. The MEG data suggested additional frontal sources. An equivalent dipole model for the generators of MEG signals indicated activity in MT+, starting at 130 ms and peaking at 170 ms after the reversal of the direction of motion, and then again at approximately 260 ms. Frontal activity began 0-20 ms later than in MT+, and peaked approximately 180 ms. Both pSTS and FEF+ showed long-duration activity continuing over the latency range of 200-400 ms. MEG responses in the region of V3A and V1/V2 were relatively small, and peaked at longer latencies than the initial peak in MT+. These data revealed characteristic patterns of activity in this cortical network for processing sudden changes in the direction of visual motion.

Parieto-occipital approximately 10 Hz activity reflects anticipatory state of visual attention mechanisms.

High-density eeg recordings revealed sensory specific modulation of anticipatory parieto-occipital approximately 10 Hz oscillatory activity when visually presented word cues instructed subjects in an intermodal selective attention paradigm. Cueing attention to the auditory features of imminent compound audio-visual stimuli resulted in significantly higher approximately 10 Hz amplitude in the period preceding onset of this stimulus than when attention was cued to the visual features. We propose that this parieto-occipital approximately 10 Hz activity reflects a disengaged visual attentional system in preparation for anticipated auditory input that is attentionally more relevant. Conversely, lower approximately 10 Hz activity during the attend-visual condition may reflect active engagement of parieto-occipital areas in the anticipatory period. These results support models implicating parieto-occipital areas in the directing and maintenance of visual attention.

Dynamic neuroimaging of brain function.

To fully characterize the brain processes underlying sensorimotor and cognitive function, the spatial distribution of active regions, their interconnected regions must be measured. We describe methods for imaging brain sources from surface-recorded EEG and magnetoencephalographic data, called electromagnetic source imaging (EMSI). EMSI provides brain source locations within the common framework of magnetic resonance (MR) images of brain anatomy. This allows integration of data from other functional brain imaging methods, like positron emission tomography and functional MR imaging, which can improve the accuracy of EMSI localization. EMSI also provides submillisecond temporal resolution of the dynamic processes within brain systems. Examples are given of applications to visual perceptual and attentional studies.

A test of brain electrical source analysis (BESA): a simulation study.

The present report summarizes the results of a simulation study on the accuracy of Scherg's implementation of spatio-temporal analysis (BESA) for estimating the parameters (wave shape, location, orientation) of the intracranial sources of event-related brain potentials (ERPs) recorded from the scalp. In view of the subjective factors that might influence a solution, 10 subjects, ranging from those with much experience with ERPs and extensive background in the use of BESA to those with little experience with BESA and/or no knowledge of ERPs, independently analyzed a set of simulated somatosensory ERP data. The simulation contained wave forms from 32 electrode sites generated by a combination of 10 dipole sources. The primary question was how faithfully the different subjects would depict the source wave shapes, locations and orientations. Based on the 9 subjects who were familiar with ERPs, the grand-average cross-correlation coefficient between subjects' estimated and actual source wave shapes was 0.89 (standard deviation (S.D.) = 0.17). The grand-average location error, based upon a head diameter of 17 cm, was 1.4 cm (S.D. = 1.0 cm). The grand-average orientation error was 24.4 degrees (S.D. = 20 degrees).

ERP amplitude and scalp distribution to target and novel events: effects of temporal order in young, middle-aged and older adults.

Event-related brain potentials (ERPs) were recorded from young (mean age = 24.1), middle-aged (48.7) and older (69.7) adults during a version of the oddball paradigm, in which 48 unique, unexpected novel stimuli were interspersed with equally rare instructed targets. As older relative to younger adults are thought to differ in their ability to inhibit the processing of task irrelevant information, we expected, based on previous work, that novel stimuli would retain their 'novelty' longer in older than in younger adults. To assess this, P3 amplitude and scalp topography elicited by novels and targets were analyzed as a function of stimulus number (n = 6) within the block and as a function of block number (n = 4). The results were in line with the prediction: While the younger adults' P3 scalp distribution shifted from a relatively more frontal to a relatively more posterior focus as a function of novel number within the block, this was not evident in the scalp topographies of the older adults. Coupled with the older adults' elevated false alarm rates to novel stimuli, the data are consistent with a change in frontal lobe function with increases in age.

Microcircuitry of posterior cingulate cortex in vitro: electrophysiology and laminar analysis using the current source density method.

We used current source density (CSD) analysis of a laminar profile of subicular stimulus-evoked field potentials recorded in cortical slices in vitro to characterize the interlaminar microcircuitry of posterior cingulate cortex. Neuroanatomic and electrophysiologic data indicate that subiculocingulate tract (SCT) afferents monosynaptically excite apical dendrites of deep laminae (V-VI) neurons, evoking pure EPSPs, while superficial laminae (II/III-IV) neurons are driven polysynaptically, evoking a mixture of longer latency EPSPs and IPSPs. Consistent with this model, CSD analysis of field potential laminar profiles supports the conclusion that activation of excitatory subicular afferent terminal fields in superficial laminae of cingulate cortex elicits primary monosynaptic activation of apical dendrites of deep lamina (V-VI) pyramids. Subsequent EPSP propagation to the somata of these pyramids generated synchronous action potential discharges which appeared to elicit delayed polysynaptic activation of superficial laminae pyramids and interneurons. Latency differences between SCT-stimulus-evoked EPSPs and action potentials in superficial and deep laminae were minimized by stimulus train frequencies of 5-8 Hz, indicating that the proposed microcircuitry can show functional tuning at frequencies characteristic of hippocampal neuronal activity (theta). Such tuning suggests that hippocampal output activity frequency and phase locked to theta rhythm will be preferentially gated through cingulate cortex.

Multiple brain systems generating the rat auditory evoked potential. I. Characterization of the auditory cortex response.

The objectives of this study were to characterize the auditory cortex response in the rat and to examine its contributions to the auditory evoked potentials (AEPs) recorded from the dorsal and lateral skull. This was accomplished by simultaneously recording AEPs from the cortical surface and from skull screw electrodes in anesthetized animals. The initial positive-negative response (P17-N32) was largely restricted to the cortical region corresponding to area 41. More detailed examination of the AEP mapping revealed multiple subcomponents (P9, P14, P17, P19) underlying the initial positivity, with differing topographies. Stimulus-response properties further dissociated the multiple positive subcomponents. Reversible local neurochemical suppression confirmed the auditory cortical origin of these AEPs. The auditory cortex-generated AEPs were refractory to barbiturate anesthesia which eliminated all dorsal skull AEPs, indicating that primary auditory cortical AEPs do not make a significant contribution to the dorsal skull-recorded ('vertex') AEPs. The findings raise issues regarding multiple parallel auditory processing systems and their associated AEPs.

Multiple brain systems generating the rat auditory evoked potential. II. Dissociation of auditory cortex and non-lemniscal generator systems.

This study addressed the issue of multiple parallel auditory processing systems and their relationship to the skull-recorded auditory evoked potentials (AEPs) in the unanesthetized, unrestrained rat. In the preceding paper (Brain Res., 602 (1993) 240-250) it has been shown that auditory cortex activity does not contribute significantly to the vertex maximal AEPs recorded from the dorsal skull of the rat. In the present study, mapping of the AEP skull distribution revealed two sets of components: one set maximal at the dorsal skull vertex, and another set at the lateral skull), but not the early (P7-P11, N15) lateral skull components generated in auditory cortex. Bilateral auditory cortex ablation eliminated the lateral skull maximal AEP components, but not the dorsal skull maximal components. These findings support extensive parallel processing of auditory inputs (reflected by the dorsal AEPs) in the absence of primary auditory cortex. Ablation of primary auditory cortex did result in a modulation of the dorsal skull AEPs, indicative of an interaction between the geniculocortical system and the parallel system which generates the dorsal AEPs.

Cellular generators of the cortical auditory evoked potential initial component.

Cellular generators of the initial cortical auditory evoked potential (AEP) component were determined by analyzing laminar profiles of click-evoked AEPs, current source density, and multiple unit activity (MUA) in primary auditory cortex of awake monkeys. The initial AEP component is a surface-negative wave, N8, that peaks at 8-9 msec and inverts in polarity below lamina 4. N8 is generated by a lamina 4 current sink and a deeper current source. Simultaneous MUA is present from lower lamina 3 to the subjacent white matter. Findings indicate that thalamocortical afferents are a generator of N8 and support a role for lamina 4 stellate cells. Relationships to the human AEP are discussed.

Generators of middle- and long-latency auditory evoked potentials: implications from studies of patients with bitemporal lesions.

We recorded middle- and long-latency auditory evoked potentials (AEPs) in 5 patients (ages 39-72 years) with bilateral lesions of the superior temporal plane. Reconstructions of CT sections revealed that primary auditory cortex had been damaged bilaterally in four of the patients, while in the fifth an extensive left hemisphere lesion included primary auditory cortex while a right hemisphere lesion had damaged anterior auditory association areas but spared primary auditory cortex. Normal middle-latency AEPs (MAEPs) were recorded at the vertex electrode in all of the patients. In 3 of the 5 patients, MAEPs also showed normal coronal scalp distributions and were comparable in amplitude following stimulation of either ear. Two patients showed abnormalities. In one case, Na (latency 17 msec)-Pa (latency 30 msec) amplitudes were reduced over both hemispheres following stimulation of the ear contralateral to the more extensive lesion. In another, with both subcortical and cortical involvement, the Pa was abolished over the hemisphere with the more extensive lesion. Long-latency AEPs were normal in 2 patients whose lesions were largely confined to the superior temporal plane. In 2 patients with lesions extending into the inferior parietal lobe, N1s were abolished bilaterally. In the fifth patient, the N1 showed a slight reduction over the hemisphere with the more extensive lesion. Middle- and long-latency AEPs were differentially affected by some lesions. For example, patients with absent N1s could produce normal Pas. A review of these results and those of previous studies of bitemporal patients suggests that abnormalities in middle- and long-latency AEPs do not necessarily reflect damage to primary auditory cortex per se, but rather the degree of damage to adjacent areas. Abnormalities in MAEPs are associated with subcortical lesions, or cortical lesions extensive enough to denervate thalamic projection nuclei. Abnormalities in the long-latency N1 reflect lesion extension into the multi-modal areas of the inferior parietal lobule. This area appears to exert a critical modulatory influence over N1 generators outside of the superior temporal plane.

Altered peripheral and brainstem auditory function in aged rats.

A technique for conducting free-field brainstem auditory evoked potential (BAEP) audiometry in unanesthetized, unrestrained rats revealed a non-recruiting 18 dB elevation of click threshold in aged rats. BAEPs were first recorded in young and aged rats to clicks of equal intensity (80 dB SPL). Compared to the young group, aged animals exhibited longer wave I and wave IV latencies with no difference seen in the I-IV central conduction time. The prominent negative wave (No) following wave IV was also delayed and the I-No and IV-No conduction times increased in the aged group. When BAEPs were recorded to clicks with intensities adjusted to 35 dB above individual threshold, no differences in wave I or wave IV latencies or in the I-IV central conduction time were found between groups. However, the No component was delayed and the I-No and IV-No conduction times remained prolonged in the aged group. The results suggest that in addition to changes in peripheral auditory structures, changes in the rostral auditory brainstem accompany age-related hearing loss in rats.

Surface auditory evoked potentials in the unrestrained rat: component definition.

Auditory evoked potentials (AEPs) to click and pure tone stimuli were recorded in unrestrained, unanesthetized rats. The middle latency rat AEPs (N17, P23, N38) had midline scalp distributions similar to human MAEPs and were recorded to within 15 dB above BAEP threshold. In contrast to human MAEPs, rat MAEPs were decreased in amplitude at high stimulation rates and only the N17 component was unaltered by slow wave sleep. The longer latency N50, N80 and P130 components had several response properties comparable to human N100-P200 vertex potentials. These included restricted midline fronto-central scalp distributions, progressive increases in amplitude at ISIs up to 4-8 sec and marked attenuation during slow wave sleep. The frequency sensitivity of the rat AEP revealed a decreased response to pure tones below 4 kHz but robust responses for stimuli up to at least 45 kHz. There was a notch in the rat audiogram with decremented component amplitudes to pure tone stimuli centered at 35 kHz. When equated for intensity, click and pure tone stimuli in the range of the rats maximal audiometric sensitivity (8-20 kHz) generated comparable AEP components. These results provide normative data on rat surface recorded AEPs. It is suggested that these surface recorded rat AEPs are generated by subcortical neural systems involved in the detection of auditory transients.

Orienting attention within visual fields: how efficient is interhemispheric transfer?

Five experiments are reported examining the effect of attentional orienting on lexical decisions within visual half-fields. In Experiment 1, following baseline performance, subjects were instructed to improve performance to the right or left of the fixation point. In Experiment 2, trials were run in blocks with all items to one side of the fixation point. In Experiment 3, completely valid position indicators as to the location of the next item to be shown were presented prior to the stimulus item. In Experiment 4, to examine practice effects, no instructions or cuing were given to subjects. In Experiment 5, subjects were urged to improve performance, but with no instructions as to location. As a summary of our results, it can be stated that (a) consistent visual field differences in lexical decision performance are present, even when subjects were informed, prior to viewing, of the spatial location of the next stimulus item. (b) Lexical decision information initially input to one cerebral hemisphere is primarily processed in that hemisphere. Interhemispheric transfer of this type of language information seems to be done primarily as the end product of a cognitive process.

The eyes have it: exposure times and saccadic movements in visual half-field experiments.

Several tachistoscopic visual half-field experiments using exposure times in excess of 150 msec have been reported and arguments have been put forth justifying this procedure. An experiment was done investigating visual field accuracy under conditions where eye movement was allowed, following parafoveal exposure. Two control experiments were done to evaluate the viewing conditions. When eye movement is permitted, accuracy in both visual fields reaches 100%. It is concluded that visual field differences found with exposure times greater than 150 msec are due to the active cooperation of the subjects and not due to the justifications advanced by experimenters using long exposure times.