Deciding where to attend: Large-scale network mechanisms underlying attention and intention revealed by graph-theoretic analysis.

The neural mechanisms by which intentions are transformed into actions remain poorly understood. We investigated the network mechanisms underlying spontaneous voluntary decisions about where to focus visual-spatial attention (willed attention). Graph-theoretic analysis of two independent datasets revealed that regions activated during willed attention form a set of functionally-distinct networks corresponding to the frontoparietal network, the cingulo-opercular network, and the dorsal attention network. Contrasting willed attention with instructed attention (where attention is directed by external cues), we observed that the dorsal anterior cingulate cortex was allied with the dorsal attention network in instructed attention, but shifted connectivity during willed attention to interact with the cingulo-opercular network, which then mediated communications between the frontoparietal network and the dorsal attention network. Behaviorally, greater connectivity in network hubs, including the dorsolateral prefrontal cortex, the dorsal anterior cingulate cortex, and the inferior parietal lobule, was associated with faster reaction times. These results, shown to be consistent across the two independent datasets, uncover the dynamic organization of functionally-distinct networks engaged to support intentional acts.

Normal aging selectively diminishes alpha lateralization in visual spatial attention.

EEG studies of cue-induced visual alpha power (8-13 Hz) lateralization have been conducted on young adults without examining differences that may develop as a consequence of normal aging. Here, we examined age-related differences in spatial attention by comparing healthy older and younger adults. Our key finding is that cue-induced alpha power lateralization was observed in younger, but not older adults, even though both groups exhibited classic event-related potential signatures of spatial orienting. Specifically, both younger and older adults showed significant early directing-attention negativity (EDAN), anterior directing-attention negativity (ADAN), late directing-attention positivity (LDAP) and contingent negative variation (CNV). Furthermore, target-evoked sensory components were enhanced for attended relative to unattended targets in both younger and older groups. This pattern of results suggests that although older adults can successfully allocate spatial attention, they do so without the lateralization of alpha power that is commonly observed in younger adults. Taken together, our findings demonstrate that younger and older adults might engage different neural mechanisms for attentional orienting, and that alpha power lateralization during visual spatial attention is a phenomenon that diminishes during normal aging.

The neural correlates of volitional attention: A combined fMRI and ERP study.

Studies of visual-spatial attention typically use instructional cues to direct attention to a relevant location, but in everyday vision, attention is often focused volitionally, in the absence of external signals. Although investigations of cued attention comprise hundreds of behavioral and physiological studies, remarkably few studies of voluntary attention have addressed the challenging question of how spatial attention is initiated and controlled in the absence of external instructions, which we refer to as willed attention. To explore this question, we employed a trial-by-trial spatial attention task using electroencephalography and functional magnetic resonance imaging (fMRI). The fMRI results reveal a unique network of brain regions for willed attention that includes the anterior cingulate cortex, left middle frontal gyrus (MFG), and the left and right anterior insula (AI). We also observed two event-related potentials (ERPs) associated with willed attention; one with a frontal distribution occurring 250-350 ms postdecision cue onset (EWAC: Early Willed Attention Component), and another occurring between 400 and 800 ms postdecision-cue onset (WAC: Willed Attention Component). In addition, each ERP component uniquely correlated across subjects with different willed attention-specific sites of BOLD activation. The EWAC was correlated with the willed attention-specific left AI and left MFG activations and the later WAC was correlated only with left AI. These results offer a comprehensive and novel view of the electrophysiological and anatomical profile of willed attention and further illustrate the relationship between scalp-recorded ERPs and the BOLD response.

Spontaneous neural fluctuations predict decisions to attend.

Ongoing variability in neural signaling is an intrinsic property of the brain. Often this variability is considered to be noise and ignored. However, an alternative view is that this variability is fundamental to perception and cognition and may be particularly important in decision-making. Here, we show that a momentary measure of occipital alpha-band power (8-13 Hz) predicts choices about where human participants will focus spatial attention on a trial-by-trial basis. This finding provides evidence for a mechanistic account of decision-making by demonstrating that ongoing neural activity biases voluntary decisions about where to attend within a given moment.

The semantic priming project.

Speeded naming and lexical decision data for 1,661 target words following related and unrelated primes were collected from 768 subjects across four different universities. These behavioral measures have been integrated with demographic information for each subject and descriptive characteristics for every item. Subjects also completed portions of the Woodcock-Johnson reading battery, three attentional control tasks, and a circadian rhythm measure. These data are available at a user-friendly Internet-based repository ( http://spp.montana.edu ). This Web site includes a search engine designed to generate lists of prime-target pairs with specific characteristics (e.g., length, frequency, associative strength, latent semantic similarity, priming effect in standardized and raw reaction times). We illustrate the types of questions that can be addressed via the Semantic Priming Project. These data represent the largest behavioral database on semantic priming and are available to researchers to aid in selecting stimuli, testing theories, and reducing potential confounds in their studies.

Attention without Constraint: Alpha Lateralization in Uncued Willed Attention.

Studies of voluntary visual spatial attention have used attention-directing cues, such as arrows, to induce or instruct observers to focus selective attention on relevant locations in visual space to detect or discriminate subsequent target stimuli. In everyday vision, however, voluntary attention is influenced by a host of factors, most of which are quite different from the laboratory paradigms that use attention-directing cues. These factors include priming, experience, reward, meaning, motivations, and high-level behavioral goals. Attention that is endogenously directed in the absence of external attention-directing cues has been referred to as "self-initiated attention" or, as in our prior work, as "willed attention" where volunteers decide where to attend in response to a prompt to do so. Here, we used a novel paradigm that eliminated external influences (i.e., attention-directing cues and prompts) about where and/or when spatial attention should be directed. Using machine learning decoding methods, we showed that the well known lateralization of EEG alpha power during spatial attention was also present during purely self-generated attention. By eliminating explicit cues or prompts that affect the allocation of voluntary attention, this work advances our understanding of the neural correlates of attentional control and provides steps toward the development of EEG-based brain-computer interfaces that tap into human intentions.

The neural markers of an imminent failure of response inhibition.

In his novel Ulysses, James Joyce wrote that mistakes are the "…portals of discovery". The present study investigated the pre-stimulus oscillatory EEG signatures of selective attention and motor preparation that predicted failures of overt response inhibition. We employed a trial-by-trial spatial cueing task using a go/no-go response paradigm with bilateral target stimuli. Subjects were required to covertly attend to the spatial location cued on each trial and respond to most of the number targets (go trials) at that location while withholding responses for one designated number (no-go trials). We analyzed the post-cue/pre-target spectral patterns comparing no-go trials in which a response occurred in error (False Alarms, FA) with trials in which participants correctly withheld a response (Correct Rejections, CR). We found that cue-induced occipital alpha (8-12 Hz) lateralization and inter-frequency anti-correlations between the motor beta (18-24 Hz) and pre-frontal theta (3-5 Hz) bands each independently predicted subsequent failures of response inhibition. Based on these findings, we infer that independent perceptual and motor mechanisms operate in parallel to contribute to failures of response inhibition.

Gating by inhibition during top-down control of willed attention.

In natural settings, the control of attention may be influenced both by external information as well as internal decision-making processes driven by intent (e.g. free will). In past studies of spatial attention, we and others have developed experimental paradigms that permit individuals to choose where to direct their attention on a trial-by-trial basis in the absence of instructive external cues - we term this willed attention. Here we investigate the electrophysiological correlates of willed attention by recording EEG activity when subjects decided to focus covert attention on one of two lateralized target locations versus when they decided to maintain attention at fixation. Independent of the direction of attention, decisions to attend, relative to decisions not to attend, resulted in significant increases in both frontal theta (4-7 Hz) power and central alpha (8-13 Hz) power. We found that focusing spatial attention, as indexed by occipital alpha lateralization was predicted across subjects by the decision-related alpha increases over central scalp regions, but not changes in frontal theta power. This finding is interpreted in terms of the Gating by Inhibition model, where the central alpha EEG signals reflect cortical inhibition of decision processes that lead to the expression of willed attention.

Variability in response criteria affects estimates of conscious identification and unconscious semantic priming.

Three experiments examined the role of response criteria in a masked semantic priming paradigm using an exclusion task. Experiment 1 used on-line prime-report ("report the prime if you saw it") and exclusion instructions in which participants were told to avoid completing a word stem (e.g. mo-) with a word related to a prime (e.g. cash) flashed for 0, 38 or 212ms. Semantic priming (i.e. exclusion failure) was significant in the items analysis, but was moderated by peoples' ability to report the prime in the participant analysis. Prime-report thresholds in Experiment 2 were made more liberal by instructing participants to guess on every trial. Prime-report increased from Experiment 1 as exclusion failures were eliminated. Experiment 3 clarified the relationship between awareness and prime identification using an on-line measure of confidence and different liberal prime report instructions. The current findings suggest that the ability to act upon (via exclusion performance) and report information in a masked prime is determined by a variable response criterion, which can be manipulated as an independent variable.

Effects of strategy on visual working memory capacity.

Substantial evidence suggests that individual differences in estimates of working memory capacity reflect differences in how effectively people use their intrinsic storage capacity. This suggests that estimated capacity could be increased by instructions that encourage more effective encoding strategies. The present study tested this by giving different participants explicit strategy instructions in a change detection task. Compared to a condition in which participants were simply told to do their best, we found that estimated capacity was increased for participants who were instructed to remember the entire visual display, even at set sizes beyond their capacity. However, no increase in estimated capacity was found for a group that was told to focus on a subset of the items in supracapacity arrays. This finding confirms the hypothesis that encoding strategies may influence visual working memory performance, and it is contrary to the hypothesis that the optimal strategy is to filter out any items beyond the storage capacity.

Age-related spatiotemporal reorganization during response inhibition.

As a key high-level cognitive function in human beings, response inhibition is crucial for adaptive behavior. Previous neuroimaging studies have shown that older individuals exhibit greater neural activation than younger individuals during response inhibition tasks. This finding has been interpreted within a neural compensation framework, in which additional neural resources are recruited in response to age-related cognitive decline. Although this interpretation has received empirical support, the precise event-related temporal course of this age-related compensatory neural response remains unexplored. In the present study, we conducted source analysis on inhibition-related ERP components (i.e., N2 and P3) that were recorded while healthy younger and older adults participated in a visual Go/NoGo task. We found that older adults showed increased source current densities of the N2 and P3 components than younger adults, which support previous hemodynamic findings. Further, such age-related differences in neural activation were successfully separated between the N2 and P3 periods by source localization analysis. Interestingly, the increased activations in older adults were primarily localized to the right precentral and postcentral gyri during the N2 period, which shifted to the right dorsolateral prefrontal cortex and the right inferior frontal gyrus during the P3 period. Taken together, our results clearly illustrate the spatiotemporal dynamics of age-related functional brain reorganization, and further specify the exact temporal course at the millisecond scale by which age-related compensatory neural responses occur during response inhibition.

The spotlight of attention illuminates failed feature-based expectancies.

A well-replicated finding is that visual stimuli presented at an attended location are afforded a processing benefit in the form of speeded reaction times and increased accuracy (Mangun, ; Posner,). This effect has been described using a spotlight metaphor, in which all stimuli within the focus of spatial attention receive facilitated processing, irrespective of other stimulus parameters. However, the spotlight metaphor has been brought into question by a series of combined expectancy studies that demonstrated that the behavioral benefits of spatial attention are contingent on secondary feature-based expectancies (Kingstone,). The present work used an event-related potential (ERP) approach to reveal that the early neural signature of the spotlight of spatial attention is not sensitive to the validity of secondary feature-based expectancies.

Individual working memory capacity is uniquely correlated with feature-based attention when combined with spatial attention.

A growing literature suggests that working memory and attention are closely related constructs. Both involve the selection of task-relevant information, and both are characterized by capacity limits. Furthermore, studies using a variety of methodological approaches have demonstrated convergent working memory and attention-related processing at the individual, neural and behavioral level. Given the varieties of both constructs, the specific kinds of attention and WM must be considered. We find that individuals' working memory capacity (WMC) uniquely interacts with feature-based attention when combined with spatial attention in a cuing paradigm (Posner, 1980). Our findings suggest a positive correlation between WM and feature-based attention only within the spotlight of spatial attention. This finding lends support to the controlled attention view of working memory by demonstrating that integrated feature-based expectancies are uniquely correlated with individual performance on a working memory task.