Oscillations in the alpha band (9-12 Hz) increase with memory load during retention in a short-term memory task

To study the role of brain oscillations in working memory, we recorded the scalp electroencephalogram (EEG) during the retention interval of a modified Sternberg task. A power spectral analysis of the EEG during the retention interval revealed a clear peak at 9-12 Hz, a frequency in the alpha band (8-13 Hz). In apparent conflict with previous ideas according to which alpha band oscillations represent brain "idling", we found that the alpha peak systematically increased with the number of items held in working memory. The enhancement was prominent over the posterior and bilateral central regions. The enhancement over posterior regions is most likely explained by the well known alpha rhythm produced close to the parietal-occipital fissure, whereas the lateral enhancement could be explained by sources in somato-motor cortex. A time-frequency analysis revealed that the enhancement was present throughout the last 2.5 s of the 2.8 s retention interval and that alpha power rapidly diminished following the probe. The load dependence and the tight temporal regulation of alpha provide strong evidence that the alpha generating system is directly or indirectly linked to the circuits responsible for working memory. Although a clear peak in the theta band (5-8 Hz) was only detectable in one subject, other lines of evidence indicate that theta occurs and also has a role in working memory. Hypotheses concerning the role of alpha band activity in working memory are discussed.     

High-resolution EEG mapping of cortical activation related to working memory: effects of task difficulty, type of processing, and practice

Changes in cortical activity during working memory tasks were examined with electroencephalograms (EEGs) sampled from 115 channels and spatially sharpened with magnetic resonance imaging (MRI)-based finite element deblurring. Eight subjects performed tasks requiring comparison of each stimulus to a preceding one on verbal or spatial attributes. A frontal midline theta rhythm increased in magnitude with increased memory load. Dipole models localized this signal to the region of the anterior cingulate cortex. A slow (low-frequency), parietocentral, alpha signal decreased with increased working memory load. These signals were insensitive to the type of stimulus attribute being processed. A faster (higher-frequency), occipitoparietal, alpha signal was relatively attenuated in the spatial version of the task, especially over the posterior right hemisphere. Theta and alpha signals increased, and overt performance improved, after practice on the tasks. Increases in theta with both increased task difficulty and with practice suggests that focusing attention required more effort after an extended test session. Decreased alpha in the difficult tasks indicates that this signal is inversely related to the amount of cortical resources allocated to task performance. Practice-related increases in alpha suggest that fewer cortical resources are required after skill development. These results serve: (i) to dissociate the effects of task difficulty and practice; (ii) to differentiate the involvement of posterior cortex in spatial versus verbal tasks; (iii) to localize frontal midline theta to the anteromedial cortex; and (iv) to demonstrate the feasibility of using anatomical MRIs to remove the blurring effect of the skull and scalp from the ongoing EEG. The results are discussed with respect to those obtained in a prior study of transient evoked potentials during working memory.      

Influence of changes in arterial carbon dioxide tension on the electroencephalogram and posterior tibial nerve somatosensory cortical evoked potentials during alfentanil/nitrous oxide anesthesia

The effects of variation of arterial CO2 tension (PaCO2) on the electroencephalogram (EEG) and posterior tibial nerve somatosensory cortical evoked potentials (PTN-SCEP) during opioid/N2O anesthesia have not been well documented. We studied the effects of hypocapnia (PaCO2 approximately 23 mmHg) and hypercapnia (PaCO2 approximately 50 mmHg) during steady-state alfentanil/N2O anesthesia in 16 patients. EEG and PTN-SCEP were recorded continuously, while PaCO2 was altered in 15-min intervals by varying the inspired CO2 concentration. Hypocapnia caused significant increases in power in the delta, theta, and beta bands (P less than 0.01), with the greatest increase observed in the alpha band. Relative power increased in the alpha band but remained unchanged in the delta, theta, and beta bands. Median frequency and 95% spectral edge frequency were unaltered during hypocapnia. In contrast, hypercapnia caused a significant decrease of power in the alpha and beta bands, whereas delta and theta power remained unchanged. This was reflected in a significant decrease of the 95% spectral edge frequency, from 8.9 (6.7-11.6) to 7.0 (5.6-8.6) Hz. All EEG parameters returned to normal upon restoration of normocapnia. There was a significant negative correlation between power in the alpha band and end-tidal CO2 in all patients (r = 0.47 to -0.89). PTN-SCEP latencies and amplitudes were not significantly different from control values during hypocapnia and hypercapnia. It is concluded that variations in PaCO2 within the limits 20-50 mmHg produce substantial changes in the EEG power spectrum, especially in the alpha band (8-12 Hz), but do not alter PTN-SCEP.     

Electroencephalographic mapping during routine clinical practice: cortical arousal during tracheal intubation?

We used quantitative analysis of the electroencephalogram (EEG) in 42 patients to assess the effect of tracheal intubation after induction of anesthesia with etomidate and sufentanil using standard clinical practice. The EEG was recorded from eight bipolar electrode derivations and Z-transformed relative to age expected normative data for relative power in the delta, theta, alpha, and beta frequency bands. Tracheal intubation resulted in classical cortical arousal, as indicated by acceleration of the EEG frequencies. Significant effects were seen in all frequency bands, most pronounced in the alpha frequency band, with the largest increase bilaterally in the fronto-temporal regions (F-values: Delta - 9.592, P < 0.001; theta - 1.691, P < 0.001; alpha - 18.439, P < 0.001; beta - 4.504, P < 0.001). Changes in alpha and delta power during induction of anesthesia were correlated with the dose of etomidate (P < 0.05). Changes in alpha after tracheal intubation were correlated at the parietooccipital brain regions to the dose of sufentanil (P < 0.05). Individual titration of the dose of etomidate and sufentanil, as during routine clinical practice, is not sufficient to block the strong noxious stimulation of tracheal intubation and results in cortical arousal. The clinical impact of this cortical wake-up phenomenon is undetermined.     

Different functional loops between cerebral cortex and the subthalmic area in Parkinson's disease

We investigate the extent to which functional circuits coupling cortical and subthalamic activity are multiple and segregated by frequency in untreated Parkinson's disease (PD). To this end, we recorded EEG and local field potentials (LFPs) from macroelectrodes inserted into the subthalamic nucleus area (SA) in nine awake patients following functional neurosurgery for PD. Patients were studied after overnight withdrawal of medication. Coherence between EEG and SA LFPs was apparent in the theta (3-7 Hz), alpha (8-13 Hz), lower beta (14-20 Hz) and upper beta (21-32 Hz) bands, although activity in the alpha and upper beta bands dominated. Theta coherence predominantly involved mesial and lateral areas, alpha and lower beta coherence the mesial and ipsilateral motor areas, and upper beta coherence the midline cortex. SA LFPs led EEG in the theta band. In contrast, EEG led the depth LFP in the lower and upper beta bands. SA LFP activity in the alpha band could either lead or lag EEG. Thus there are several functional sub-loops between the subthalamic area and cerebral cortical motor regions, distinguished by their frequency, cortical topography and temporal relationships. Tuning to distinct frequencies may provide a means of marking and segregating related processing, over and above any anatomical segregation of processing streams.     

Changes in cortical electrical activity during induction of anaesthesia with thiopental/fentanyl and tracheal intubation: a quantitative electroencephalographic analysis

Background. There are regional differences in the effects ofanaesthetics agents and perioperative stimuli on the EEG. Westudied the topography of the EEG during induction of anaesthesiaand intubation in patients receiving thiopental and fentanylto document regional electrical brain activity. Methods. EEG was recorded in 25 patients in the awake state,after pre-medication, during induction, at loss of consciousnessand after intubation. Eight bipolar recordings were made andthe relative power of the frequency bands delta, theta, alpha,and beta were used (after z-score transformation for age) tomeasure changes in regional EEG activity. Results. Noxious stimulation during tracheal intubation partiallyreversed the slowing of the EEG caused by anaesthesia. Duringinduction of anaesthesia alpha activity was most reduced intemporal and occipital regions. The most prominent EEG changesafter intubation were an increase in alpha and a decrease indelta power (P<0.001). The largest changes were in the frontaland temporal leads for alpha and in the frontal and centralleads for delta. Heart rate and arterial pressure remained constantduring intubation. Conclusions. Changes in alpha and delta power were identifiedas the most sensitive EEG measures of regional changes in electricalbrain activity during anaesthesia and noxious stimulation. Br J Anaesth 2004; 92: 33–8     

Neurophysiological measures of working memory and individual differences in cognitive ability and cognitive style

The capacity to deliberately control attention in order to hold and manipulate information in working memory is critical to higher cognitive functions. This suggests that between-subject differences in general cognitive ability might be related to observable differences in the activity of brain systems that support working memory and attention control. To test this notion, electroencephalograms were recorded from 80 healthy young adults during spatial working memory tasks. Measures of task-related neurophysiological and behavioral variables were derived from these data and compared to scores on a test battery commonly used to assess general cognitive ability (the WAIS-R). Subjects who scored high on the psychometric test also tended to respond faster in the experimental tasks without any loss of accuracy. The amplitude of the late positive component of the event-related potential was larger in high-ability subjects, and the frontal midline theta component of the EEG signal was also selectively enhanced in this group under conditions of sustained performance and high working memory load. These results suggest that subjects who scored high on the WAIS-R were better able to focus and sustain attention to task performance. Changes in the EEG alpha rhythm in response to manipulations of task practice and load were also examined and compared between frontal and parietal regions. The results indicated that high-ability subjects developed strategies that made relatively greater use of parietal regions, whereas low-ability subjects relied more exclusively on frontal regions. Other analyses indicated that hemispheric asymmetries in alpha band measures distinguish between individuals with relatively high verbal aptitude and those with relatively high nonverbal aptitude. In particular, subjects with a verbal cognitive style tended to make greater use of the left parietal region during task performance, and subjects with a nonverbal style tended to make greater use of the right parietal region. These results help clarify relationships between task-related brain activity and individual differences in cognitive ability and style.     

Oscillatory activity in human parietal and occipital cortex shows hemispheric lateralization and memory effects in a delayed double-step saccade task

We applied magnetoencephalography (MEG) to record oscillatory brain activity from human subjects engaged in planning a double-step saccade. In the experiments, subjects (n = 8) remembered the locations of 2 sequentially flashed targets (each followed by a 2-s delay), presented in either the left or right visual hemifield, and then made saccades to the 2 locations in sequence. We examined changes in spectral power in relation to target location (left or right) and memory load (one or two targets), excluding error trials based on concurrent eye tracking. During the delay period following the first target, power in the alpha (8-12 Hz) and beta (13-25 Hz) bands was significantly suppressed in the hemisphere contralateral to the target. When the second target was presented, there was a further suppression in the alpha- and beta-band power over both hemispheres. In this period, the same sensors also showed contralateral power enhancements in the gamma band (60-90 Hz), most significantly prior to the initiation of the saccades. Adaptive spatial filtering techniques localized the neural sources of the directionally selective power changes in parieto-occipital areas. These results provide further support for a topographic organization for delayed saccades in human parietal and occipital cortex.     

Pre- and poststimulus alpha rhythms are related to conscious visual perception: a high-resolution EEG study

Conscious and unconscious visuospatial processes have been related to parietooccipital cortical activation as revealed by late visual-evoked potentials. Here the working hypothesis was that a specific pattern of pre- and poststimulus theta (about 4-6 Hz) and alpha (about 6-12 Hz) rhythms is differently represented during conscious compared with unconscious visuospatial processes. Electroencephalographic (EEG) data (128 channels) were recorded in normal adults during a visuospatial task. A cue stimulus appeared at the right or left (equal probability) monitor side for a "threshold time" inducing about 50% of correct recognitions. It was followed (2 s) by visual go stimuli at spatially congruent or incongruent position with reference to the cue location. Left (right) mouse button was clicked if the go stimulus appeared at the left (right) monitor side. Then, subjects said "seen" if they had detected the cue stimulus or "not seen" if missed (self-report). Sources of theta and alpha rhythms during seen and not seen EEG epochs were estimated by low-resolution electromagnetic brain topography software. Results showed that the prestimulus "low-band" (about 6-10 Hz) alpha rhythms in frontal, parietal, and occipital areas were stronger in power in the seen than in the not seen trials. After the visual stimulation, the power of the "high-band" (about 10-12 Hz) alpha rhythms in parietal and occipital areas decreased more in the seen than in the not seen trials. The present results suggest that visuospatial consciousness covary--presumably with a facilitatory effect--with the power of both pre- and poststimulus alpha rhythms.     

Topography of Clonidine-induced Electroencephalographic Changes Evaluated by Principal Component Analysis

Background: Principal component analysis is a multivariate statistical technique to facilitate the evaluation of complex data dimensions. In this study, principle component analysis was used to reduce the large number of variables from multichannel electroencephalographic recordings to a few components describing changes of spatial brain electric activity after intravenous clonidine.

Methods: Seven healthy volunteers (age, 26 +/- 3 [SD] yr) were included in a double-blind crossover study with intravenous clonidine (1.5 and 3.0 [mu]g/kg). A spontaneous electroencephalogram was recorded by 26 leads and quantified by standard fast Fourier transformation in the [delta], [theta], [alpha], and [beta] bands. Principle component analysis derived from a correlation matrix calculated between all electroencephalographic leads (26 x 26 leads) separately within each classic frequency band. The basic application level of principle component analysis resulted in components representing clusters of electrodes positions that were differently affected by clonidine. Subjective criteria of drowsiness and anxiety were rated by visual analog scales.

Results: Topography of clonidine-induced electroencephalographic changes could be attributed to two independent spatial components in each classic frequency band, explaining at least 85% of total variance. The most prominent effects of clonidine were increases in the delta band over centroparietooiccipital areas and decreases in the alpha band over parietooccipital regions. Clonidine administration resulted in subjective drowsiness.     

梅毒晚期麻痹性痴呆患者的脑电图特征分析

目的探讨梅毒晚期麻痹性痴呆患者的脑电图特征及其诊断价值。方法选取2011年5月至2013年12月首都医科大学附属北京地坛医院神经内科收治的麻痹性痴呆患者11例和同期健康对照组11例做为研究对象。在Matlab 7.0平台上对脑电信号进行频谱分析,分别计算各导联（Fp1、Fp2、F3、F4、C3、C4、P3、P4、O1、O2、F7、F8、T3、T4、T5、T6）各频带（δ、θ、α、β、γ频带）的相对功率比、α/θ值及左右半球间的相应导联（Fp1-Fp2、F3-F4、C3-C4、P3-P4、O1-O2、F7-F8、T3-T4、T5-T6）各频带（δ、θ、α、β、γ频带）的最大相干值及其对应频率。结果在相对功率比的比较中,麻痹性痴呆患者较对照组慢波频带（δ、θ频带）升高（P分别〈0.05、〈0.01和〈0.001）,快波频带（α、β、γ频带）降低（P分别〈0.05、〈0.01和〈0.001）;在双额、双枕、右额、右颞及左顶区麻痹性痴呆组α/θ值降低（P分别〈0.05和〈0.01）;额顶区γ频带、顶区β频带、枕区α频带和颞区θ频带的最大相干值及顶区α频带,枕区β频带,颞区θ、γ、δ频带的对应频率较对照组降低（P分别〈0.05和〈0.01）。结论麻痹性痴呆患者的脑电慢波频带呈升高趋势,快波频带呈降低趋势,左右半球间相应导联频带的最大相干值及对应频率减低,可为诊断提供有力的客观依据。     

Que PASA? The posterior-anterior shift in aging

A consistent finding from functional neuroimaging studies of cognitive aging is an age-related reduction in occipital activity coupled with increased frontal activity. This posterior-anterior shift in aging (PASA) has been typically attributed to functional compensation. The present functional magnetic resonance imaging sought to 1) confirm that PASA reflects the effects of aging rather than differences in task difficulty; 2) test the compensation hypothesis; and 3) investigate whether PASA generalizes to deactivations. Young and older participants were scanned during episodic retrieval and visual perceptual tasks, and age-related changes in brain activity common to both tasks were identified. The study yielded 3 main findings. First, inconsistent with a difficulty account, the PASA pattern was found across task and confidence levels when matching performance among groups. Second, supporting the compensatory hypothesis, age-related increases in frontal activity were positively correlated with performance and negatively correlated with the age-related occipital decreases. Age-related increases and correlations with parietal activity were also found. Finally, supporting the generalizability of the PASA pattern to deactivations, aging reduced deactivations in posterior midline cortex but increased deactivations in medial frontal cortex. Taken together, these findings demonstrate the validity, function, and generalizability of PASA, as well as its importance for the cognitive neuroscience of aging.     

High-density EEG mapping during general anaesthesia with xenon and propofol: a pilot study

Anaesthetic-induced spatial inhomogeneities of the electrencephalogram(EEG) using "high density" electrode mapping have not previously been reported. We measured the scalp EEG with a dense electrode (128-channel) montage during the course of light general anaesthesia with xenon and then propofol in normal human subjects. EEG was measured during induction and recovery of general anaesthesia in five normal subjects, and we obtained analysable data from three of these subjects. EEG topographies were plotted on a realistic head surface. Scalp fields were spatially de-blurred using a realistic head model and projected onto an averaged cortical surface Both xenon and propofol elicited large increases in midline frontal theta-band EEG power. Propofol reliably elicited orbitofrontal delta activity. Xenon, but not propofol, caused large increases in delta over the posterior cortex. Increased gamma power was observed for both anaesthetic agents at midline electrodes over the posterior cortex, but not anteriorly. Anaesthesia-induced delta and theta waves were differentially distributed along the anterior-posterior axis of the brain in a manner that corresponds well to the anatomy of putative neuronal generators. The distribution of anaesthetic-induced changes in fast gamma-band power seems to reflect functional differences between the posterior and anterior aspects of the cerebral cortex. These preliminary observations were consistent within our small sample, indicating that larger studies of anaesthetic effects using high-density recordings are warranted.     

Topography of clonidine-induced electroencephalographic changes evaluated by principal component analysis

BACKGROUND: Principal component analysis is a multivariate statistical technique to facilitate the evaluation of complex data dimensions. In this study, principle component analysis was used to reduce the large number of variables from multichannel electroencephalographic recordings to a few components describing changes of spatial brain electric activity after intravenous clonidine. METHODS: Seven healthy volunteers (age, 26 +/- 3 [SD] yr) were included in a double-blind crossover study with intravenous clonidine (1.5 and 3.0 microg/kg). A spontaneous electroencephalogram was recorded by 26 leads and quantified by standard fast Fourier transformation in the delta, theta, alpha, and beta bands. Principle component analysis derived from a correlation matrix calculated between all electroencephalographic leads (26 x 26 leads) separately within each classic frequency band. The basic application level of principle component analysis resulted in components representing clusters of electrodes positions that were differently affected by clonidine. Subjective criteria of drowsiness and anxiety were rated by visual analog scales. RESULTS: Topography of clonidine-induced electroencephalographic changes could be attributed to two independent spatial components in each classic frequency band, explaining at least 85% of total variance. The most prominent effects of clonidine were increases in the delta band over centroparietooiccipital areas and decreases in the alpha band over parietooccipital regions. Clonidine administration resulted in subjective drowsiness. CONCLUSIONS: Data from the current study supported the fact that spatial principle component analysis is a useful multivariate statistical procedure to evaluate significant signal changes from multichannel electroencephalographic recordings and to describe the topography of the effects. The clonidine-related changes seen here were most probably results of its sedative effects.     

Spectrotemporal analysis of evoked and induced electroencephalographic responses in primary auditory cortex (A1) of the awake monkey

Electroencephalography is increasingly being used to probe the functional organization of auditory cortex. Modulation of the electroencephalographic (EEG) signal by tones was examined in primary auditory cortex (A1) of awake monkeys. EEG data were measured at 4 laminar depths defined by current source density profiles evoked by best frequency (BF) tones. Midlaminar multiunit activity was used to define the tuning characteristics of A1 sites. Presentation of BF tones increased EEG power across the range of frequencies examined (4-290 Hz), with maximal effects evident within the first 100 ms after stimulus onset. The largest relative increases in EEG power generally occurred at very high gamma frequency bands (130-210 Hz). Increases in EEG power for frequencies less than 70 Hz primarily represented changes in phase-locked activity, whereas increases at higher frequencies primarily represented changes in non-phase-locked activity. Power increases in higher gamma bands were better correlated with the A1 tonotopic organization than power increases in lower frequency bands. Results were similar across the 4 laminar depths examined. These findings highlight the value of examining high-frequency EEG components in exploring the functional organization of auditory cortex and may enhance interpretation of related studies in humans.     

Attentional modulation of SSVEP power depends on the network tagged by the flicker frequency

Modulation of the steady-state visual evoked potential (SSVEP) by attention was studied in detail using 15 'tag' frequencies in the range of 2.5-20 Hz. The stimuli were two series of random disc search arrays superimposed on two concentric color-marked annuli respectively. Two series of arrays were updated independently; one updated at one fixed frequency (flicker) and the other updated randomly according to a white noise distribution (random broadband flicker, rbbf). On each trial, the observer was instructed to attend one annulus and to detect a target (a triangle) that occasionally appeared in a random disc array in the attended annulus. The SSVEP results show that the choice of flicker frequency selects which cortical network synchronizes to the flicker two distinct cortical networks showed different effects of attention. SSVEP power and the effects of attention on SSVEP power strongly depend on both flicker frequency and radial position of rbbf annulus. At flicker frequencies in the delta band (2-4 Hz), and in the upper alpha band (10-11 Hz), an occipital-frontal network appears to phase-lock to the flicker when attending to the flicker, increasing the magnitude of the SSVEP. At flicker frequencies in the lower alpha band (8-10 Hz), a global response to a peripheral flickering stimulus, that includes parietal cortex and posterior frontal cortex, has higher amplitude when attention is directed away from the flickering peripheral stimulus and towards a competing rbbf stimulus in the fovea. Increases in SSVEP power when attention is directed to peripheral flicker are always associated with increases in phase locking. By contrast, at frequencies in the lower alpha band, increases in SSVEP power when attention is directed away from the flicker and towards foveal stimuli are not associated with changes in phase-locking. Thus, whether attention to a flicker stimulus increases or decreases SSVEP amplitude and phase locking depends on which of two cortical networks, which have distinct spatial and dynamic properties, is selected by the flicker frequency.     

Oscillatory EEG correlates of episodic trace decay

Recent studies suggest that human theta oscillations appear to be functionally associated with memory processes. It is less clear, however, to what type of memory sub-processes theta is related. Using a continuous word recognition task with different repetition lags, we investigate whether theta reflects the strength of an episodic memory trace or general processing demands, such as task difficulty. The results favor the episodic trace decay hypothesis and show that during the access of an episodic trace in a time window of approximately 200-400 ms, theta power decreases with increasing lag (between the first and second presentation of an item). LORETA source localization of this early theta lag effect indicates that parietal regions are involved in episodic trace processing, whereas right frontal regions may guide the process of retrieval. We conclude that episodic encoding can be characterized by two different stages: traces are first processed at parietal sites at approximately 300 ms, then further processing takes place in regions of the medial temporal lobe at approximately 500 ms. Only the first stage is related to theta, whereas the second is reflected by a slow wave with a frequency of approximately 2.5 Hz.     

Regional differences of the human sleep electroencephalogram in response to selective slow-wave sleep deprivation

The purpose of this study was to assess the topographic changes in sleep recuperative processes in response to selective slow-wave sleep (SWS) deprivation. SWS was suppressed on two consecutive nights by means of acoustic stimulation. The electroencephalogram (EEG) power of baseline, deprivation and recovery nights was analysed in 1 Hz bins. During the SWS deprivation nights, large decreases of EEG power were found at frontopolar, central and parietal derivations encompassing the delta, theta and alpha range, while only slow delta (0.5-2 Hz) was affected at the frontal derivation. Recovery sleep was characterized by a generalized increase of power during non-REM sleep encompassing the delta, theta and alpha bands, with a clear antero-posterior gradient. The coherent behaviour of different EEG bands with traditionally different electrophysiological meanings during non-REM sleep suggests that, in light of the recent advances in sleep neurophysiology, a re-examination of the functional role of EEG rhythms during sleep is needed. The 'resistance' to selective SWS deprivation of the frontal area, together with its larger increase of EEG power during recovery, may be interpreted as a sign of a greater sleep need of the frontal cortical areas, confirming that some aspects of the regulatory processes of human sleep are local in nature and may show use-dependent characteristics.     

Competition in working memory reduces frontal guidance of visual selection

Working memory (WM) representations can bias visual selection to matching stimuli in the field. WM biases can, however, be modulated by the level of cognitive load, with WM guidance reduced as memory load increases. Here, we used functional magnetic resonance imaging to distinguish between competing hypotheses for the reduction of WM guidance under load: 1) poor neural representations of memory contents under high load, 2) strategic control at high loads to direct attention away from search distracters matching the WM content, and 3) reduction of frontal top-down biasing of visual areas with increasing memory loads. We show that matching between WM contents and the visual display appeared to be well represented in visual areas under high memory loads, despite a lack of WM guidance at the behavioral level. There was little engagement of "cognitive control" areas in the prefrontal cortex during search at high loads. More importantly, WM guidance at low loads engaged a set of frontal regions in the superior and inferior ventral frontal cortex. Functional connectivity analyses revealed frontal regions working in concert with occipital areas at low memory loads, but this coupling was disrupted by increased memory load. We discuss the implications for understanding the mechanisms supporting the interplay between WM and attention.     

Retinotopy and attention in human occipital, temporal, parietal, and frontal cortex

Novel mapping stimuli composed of biological motion figures were used to study the extent and layout of multiple retinotopic regions in the entire human brain and to examine the independent manipulation of retinotopic responses by visual stimuli and by attention. A number of areas exhibited retinotopic activations, including full or partial visual field representations in occipital cortex, the precuneus, motion-sensitive temporal cortex (extending into the superior temporal sulcus), the intraparietal sulcus, and the vicinity of the frontal eye fields in frontal cortex. Early visual areas showed mainly stimulus-driven retinotopy; parietal and frontal areas were driven primarily by attention; and lateral temporal regions could be driven by both. We found clear spatial specificity of attentional modulation not just in early visual areas but also in classical attentional control areas in parietal and frontal cortex. Indeed, strong spatiotopic activity in these areas could be evoked by directed attention alone. Conversely, motion-sensitive temporal regions, while exhibiting attentional modulation, also responded significantly when attention was directed away from the retinotopic stimuli.