How do we select perceptions and actions? Human brain imaging studies

The selective nature of human perception and action implies a modulatory interaction between sensorimotor processes and attentional processes. This paper explores the use of functional imaging in humans to explore the mechanisms of perceptual selection and the fate of irrelevant stimuli that are not selected. Experiments with positron emission tomography show that two qualitatively different patterns of modulation of cerebral blood flow can be observed in experiments where non-spatial visual attention and auditory attention are manipulated. These patterns of modulation of cerebral blood flow modulation can be described as gain control and bias signal mechanisms. In visual and auditory cortex, the dominant change in cerebral blood flow associated with attention to either modality is related to a bias signal. The relation of these patterns of modulation to attentional effects that have been observed in single neurons is discussed. The existence of mechanisms for selective perception raises the more general question of whether irrelevant ignored stimuli are nevertheless perceived. Lavie's theory of attention proposes that the degree to which ignored stimuli are processed varies depending on the perceptual load of the current task. Evidence from behavioural and functional magnetic resonance imaging studies of ignored visual motion processing is presented in support of this proposal.     

Manipulating attentional asymmetry affects self-reported arousal.

This investigation examined whether manipulating participants' attention toward the left or right visual field would have consequences for mood state. Based on the premise that right parietal activity is related to the arousal dimension of emotion, it was predicted that activating the right parietal region asymmetrically would differentially affect aspects of mood related to arousal. Half of the participants in the experiment completed a leftward-biased attentional orienting task, and half completed a mirror-image rightward-biased task. Consistent with predictions, participants in the left-biased group showed greater changes in self-reported arousal. Results support the notion of bidirectional causal relations between mood and perceptual asymmetries. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Asymmetric neural control systems in human self-regulation.

Reviews the literature on the neurotransmitter substrates controlling motor readiness, showing that these substrates produce qualitative changes in the flow of information in the brain: Dopaminergic activation increases informational redundancy, whereas noradrenergic arousal facilitates orienting to novelty. Evidence that these neurotransmitter pathways are lateralized in the human brain is consistent with the left hemisphere's specialization for complex motor operations and the right hemisphere's integration of bilateral perceptual input. Principles of attentional control are suggested by the operational characteristics of neural control systems. The affective features of the activation and arousal systems are integral to their adaptive roles and may suggest how specific emotional processes dynamically regulate cognitive function. (4? p ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Mild stress increases attentional bias in social drinkers who drink to cope: A replication and extension.

The authors investigated the effects of a laboratory stressor task on rapid initial orienting and delayed disengagement aspects of attentional bias for alcohol-related cues in social drinkers (N = 58). Participants were randomly allocated to receive a stress induction or control manipulation before completing a visual probe task in which alcohol cues were presented for either 100 ms or 500 ms. Results indicated that participants who reported drinking alcohol to cope with negative affect had increased attentional bias for alcohol cues after stress induction. This effect of stress induction on attentional bias was evident when cues were presented for 100 ms and 500 ms, which suggests that stress increases both the initial orienting of attention toward and the delayed disengagement of attention from alcohol-related cues. Theoretical implications of this robust finding are discussed. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Contributions of the dopaminergic system to voluntary and automatic orienting of visuospatial attention

Visuospatial attention can be directed by voluntary or involuntary control independent of eye movement. The involvement of cortical and subcortical neural structures in this covert orienting mechanism has been studied using neuroimaging and electrophysiological techniques. This study was designed to investigate the role of the dopaminergic system in both voluntary and automatic orienting mechanisms of visuospatial attention. We recorded event-related evoked potentials (ERPs) and reaction time (RT) during a cued priming task in both patients with idiopathic Parkinson's disease (PD) and control subjects. Voluntary and automatic shifts in attention were studied by using central and peripheral cues, respectively. In the experiment using a central cue, the RT data showed that when the cue-target interval was long, PD patients' responses were delayed, and cue validity effects were reduced, whereas in the peripheral cue experiment the validity effects persisted across all trials. The ERPs demonstrated reduced sustained negativities preceding the imperative targets in both the central and peripheral cue experiments in PD patients. Furthermore, during the long cue-target interval in the central cue experiment, PD patients showed reduced attention shift-related negativities (ARNs) at the anterior scalp sites, whereas ARNs were generated widely in the peripheral cue experiment. The ERP findings were consistent with the RT data. These findings suggest that the dopaminergic system may contribute to voluntary and sustained control of visuospatial attention as well as to the neural system for response preparation, whereas automatic control of visuospatial attention is relatively independent of the dopamine system.     

The neuropsychology of 3-D space

The neuropsychological literature on 3-D spatial interactions is integrated using a model of 4 major behavioral realms: (a) peripersonal (visuomotor operations in near-body space), (b) focal extrapersonal (visual search and object recognition), (c) action extrapersonal (orienting in topographically defined space), and (d) ambient extrapersonal (orienting in earth-fixed space). Each is associated with a distinct cortical network: dorsolateral peripersonal, predominantly ventrolateral focal-extrapersonal, predominantly ventromedial action-extrapersonal, and predominantly dorsomedial ambient-extrapersonal systems. Interactions in 3-D space are also regulated neurochemically with dopaminergic and cholinergic excitation associated with extrapersonal activation and noradrenergic and serotonergic excitation associated with peripersonal activation. This model can help explain the 3-D imbalances in prominant neuropsychological disorders.     

Cholinergic influences on feature binding.

The binding problem refers to the fundamental challenge of the central nervous system to integrate sensory information registered by multiple brain regions to form a unified neural representation of a stimulus. Human behavioral, neuropsychological, and functional neuroimaging evidence suggests a fundamental role for attention in feature binding; however, its neurochemical basis is currently unknown. This study examined whether acetylcholine (ACh), a neuromodulator that has been implicated in attentional processes, plays a critical role in feature binding. Using a within-subjects pharmacological design and the cholinergic muscarinic antagonist scopolamine, the present experiments demonstrate, in a rat model, a critical role for the cortical muscarinic cholinergic system in feature binding. Specifically, ACh and the attentional resources that it supports are essential for the initial feature binding process but are not required to maintain neural representations of bound stimuli. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The functional neuroanatomy of awareness: with a focus on the role of various anatomical systems in the control of intermodal attention

This review considers a number of recent theories on the neural basis of consciousness, with particular attention to the theories of Bogen, Crick, Llinás, Newman, and Changeux. These theories allot different roles to various key brain areas, in particular the reticular and intralaminar nuclei of the thalamus and the cortex. Crick's hypothesis is that awareness is a function of reverberating corticothalamic loops and that the spotlight of intramodal attention is controlled by the reticular nucleus of the thalamus. He also proposed different mechanisms for attention and intention ("will"). The current review presents a new hypothesis, based on elements from these hypotheses, including intermodal attention and olfaction and pain, which may pose problems for Crick's original theory. This work reviews the possible role in awareness and intermodal attention and intention of the cholinergic system in the basal forebrain and the tegmentum; the reticular, the intralaminar, and the dorsomedial thalamic nuclei; the raphe and locus coeruleus; the reticular formation; the ventral striatum and extended amygdala; insula cortex, and other selected cortical, areas. Both clinical and basic research data are covered. The conclusion is reached that the brain may work by largely nonlinear parallel processing and much intramodal shifts of attention may be effected by intracortical, or multiple corticothalamic mechanisms (small local "flashlights" rather than one major "searchlight"). But this is constrained by the functional anatomy of the circuits concerned and waking "awareness" is modulated by the many "nonspecific" systems (cholinergic from the basal forebrain, noradrenergic from the locus coeruleus, dopaminergic from the substantia nigra and ventral tegmentum, and serotoninergic from the raphe). But the principal agents for intermodal attention shifts, the "searchlight," may be two key nuclei of the cholinergic system in the mesencephalon. Clinical loss of consciousness results from damage to these nuclei but not from damage to the cholinergic nucleus basalis of the basal forebrain.     

Neural substrates of latent inhibition: The switching model.

Latent inhibition (LI) refers to decrement in conditioning to a stimulus as a result of its prior nonreinforced preexposure. This robust phenomenon has been shown in classical and instrumental conditioning procedures and in many mammalian species, including humans. Development of LI reflects decreased associability of, or attention to, stimuli that predict no significant outcome. The fact that LI reflects attentional processes has become important to neuroscientists who see LI as a convenient tool for measuring the effects of drug treatments and lesions on attention. Data on brain systems studied for their involvement in LI are surveyed. These are presented in sections on noradrenergic, cholinergic, dopaminergic, serotonergic, and septo-hippocampal manipulations. It is concluded that the neural substrates of LI include the mesolimbic dopaminergic system (MDS), the mesolimbic serotonergic system (MSS), and the hippocampus. The preexposed stimulus loses its capacity to affect behavior in conditioning, even though it predicts reinforcement, because the hippocampus inhibits the switching mechanism of the nucleus accumbens via the subiculum-accumbens pathway. This hippocampal action is modulated by the MSS via its interactions with the hippocampal system or MDS, or both. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Attentional effects of single dose triazolam

1. While the effects of benzodiazepines on human memory have been extensively studied little is known about the effects of these agents on attentional processes. The authors studied the effects of a single dose of triazolam on selective visual-spatial attention using a double blind, placebo controlled design. 2. In each of 2 sessions 12 normal volunteers ingested either 0.25 mg of triazolam or placebo. Attentional performance was evaluated using two versions of the covert orienting paradigm which measured automatic (exogenous) and controlled (endogenous) aspects of attentional orienting, respectively. 3. Triazolam selectively modified performance on automatic orienting to exogenous cues. Specifically, triazolam increased the facilitation of target detection seen at shorter (150 msec) SOA's. This may indicate an increase in facilitation and a reduction in inhibition or a slowing of the time course of the biphasic attentional effect normally resulting from exogenous cuing. 4. These results indicate the importance of using experimental paradigms which effectively dissociate endogenous and exogenous mechanisms of spatial orienting in studies evaluating the effects of pharmacological agents on visual-spatial attention.     

What can be learned about attention from studying its development?

Suggests the importance of developmental research in evaluating theories of visual attention. Developmental studies can and do provide a unique vantage point from which to assess theories of attention. Three research steps are proposed: (1) theoretically-important differences and similarities between age groups are established, (2) theoretical constructs are mapped in these age differences/similarities, and (3) data are collected to examine the relation between the constructs and age. Several examples of the use of this strategy are summarized. The variable of age is shown to play a role in testing theories of Gestalt grouping, perceptual organization, spatial orienting, and attentional filtering. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Attentional bias in drug dependence: Vigilance for cigarette-related cues in smokers.

Two experiments investigated attentional biases for smoking-related cues in smokers and nonsmokers, using the visual probe task. In Experiment 1, when pictures were displayed for 500 ms, smokers who had made repeated quit attempts showed an attentional bias for smoking-related scenes. Experiment 2 replicated this finding and revealed that when pictures were presented for 2,000 ms, the smoker group as a whole showed vigilance for smoking-related cues, but nonsmokers did not. The findings from the 500-ms exposure condition suggest that initial orienting of attention to smoking cues was associated with repeated unsuccessful attempts at abstinence in smokers. Results are discussed with reference to incentive-sensitization theories of addiction and to component processes of selective attention, such as initial orienting versus maintenance. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Differential modulation of high-frequency gamma-electroencephalogram activity and sleep-wake state by noradrenaline and serotonin microinjections into the region of cholinergic basalis neurons

Several lines of evidence indicate that cholinergic basalis neurons play an important role in cortical activation. The present study was undertaken to determine the effect of noradrenergic and serotonergic modulation of the cholinergic neurons on cortical EEG activity and sleep-wake states. The neurotransmitters were injected into the region of the basalis neurons by remote control in freely moving, naturally sleeping-waking rats during the day when the rats are normally asleep the majority of the time. Effects were observed on behavior and EEG activity, including high-frequency gamma activity (30-60 Hz), which has been demonstrated to reflect behavioral and cortical arousal in the rat. Noradrenaline, which has been shown in previous in vitro studies to depolarize and excite the cholinergic cells, produced a dose-dependent increase in gamma-EEG activity, a decrease in delta activity, and an increase in waking. Serotonin, which has been found in previous in vitro studies to hyperpolarize the cholinergic neurons, produced a dose-dependent decrease in gamma-EEG activity with no significant change in amounts of wake or slow wave sleep. Both chemicals resulted in a dose-dependent decrease in paradoxical sleep. These results demonstrate that noradrenaline and serotonin exert differential modulatory effects on EEG activity through the basal forebrain, the one facilitating gamma activity and eliciting waking and the other diminishing gamma activity and not significantly affecting slow wave sleep. The results also confirm that the cholinergic basalis neurons play an important role in cortical activation and particularly in the high-frequency gamma activity that underlies cortical and behavioral arousal of the wake state.     

Attention and brain function in Alzheimer's disease: A review.

Cognitive and neuroscience studies point to a selective impairment of attentional functions in Alzheimer's disease (AD). Prominent deficits occur in the shifting and division of attention, whereas phasic arousal and focused attention to stimulus features are only minimally affected in the early stages of AD. Macroscopic processing interpretations of these results—global cognitive impairment, generalized cognitive slowing, processing resource deficit, and failure of central executive control ("dysexecutive" syndrome)—are discussed. An alternative approach is based on the identification of component attentional operations and their mediation by corticocortical and subcortical networks. This analysis suggests that attention represents the first cognitive indicator of neocortical dysfunction in early AD. Disconnection between frontal and posterior parietal areas may mediate the selective disruption of attentional functions in AD. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Dissociation of attention in learning and action: Effects of lesions of the amygdala central nucleus, medial prefrontal cortex, and posterior parietal cortex.

Many associative learning theories assert that the predictive accuracy of events affects the allocation of attention to them. More reliable predictors of future events are usually more likely to control action based on past learning, but less reliable predictors are often more likely to capture attention when new information is acquired. Previous studies showed that a circuit including the amygdala central nucleus (CEA) and the cholinergic substantia innominata/nucleus basalis magnocellularis (SI/nBM) is important for both sustained attention guiding action in a five-choice serial reaction time (5CSRT) task and for enhanced new learning about less predictive cues in a serial conditioning task. In this study, the authors found that lesions of the cholinergic afferents of the medial prefrontal cortex interfered with 5CSRT performance but not with surprise-induced enhancement of learning, whereas lesions of cholinergic afferents of posterior parietal cortex impaired the latter effects but did not affect 5CSRT performance. CEA lesions impaired performance in both tasks. These results are consistent with the view that CEA affects these distinct aspects of attention by influencing the activity of separate, specialized cortical regions via modulation of SI/nBM. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Specific changes in conditioned responding following neurotoxic damage to the posterior parietal cortex.

The central nucleus (CN) of the amygdala and basal forebrain cholinergic projections to the posterior parietal cortex (PPC) are involved in regulating changes in attentional processing of conditioned stimuli. In a previous study, lesions of the CN produced a deficit in conditioned orienting behavior (rearing on the hind legs) when a visual stimulus was paired with food. Unconditioned orienting (rearing to nonreinforced presentations of the stimulus) and conditioned food cup behavior were unaffected. The present study examined the contribution of the PPC to attentional orienting behavior. Damage to the PPC did not affect orienting behavior but produced deficits in food cup behavior. These findings help define the specific contributions of the PPC to attentional processing and associative learning. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Disconnection of the amygdala central nucleus and the substantia innominata/nucleus basalis magnocellularis disrupts performance in a sustained attention task.

The basal forebrain cholinergic system is broadly implicated in the regulation of attention. Disruptions in the function of this system produce impairments in many attentional functions, including the performance of well-learned responses under increased attentional load and the surprise-induced enhancement of learning rate. Similarly, lesions of the amygdala central nucleus (CeA) have been found to impair attentional function in some circumstances. In the present article, the effects of lesions that disconnected CeA from the cholinergic substantia innominata/nucleus basalis magnocellularis (SI/nBM) on performance are examined in a modified 5-choice serial reaction time (5CSRT) task, thought to assess selective or sustained attention. The lesions impaired performance under conditions of increased attentional load, suggesting that a circuit that includes CeA and SI/nBM regulates these aspects of attention. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Multiple mechanisms of visual-spatial attention: recent evidence from human electrophysiology

Natural visual scenes contain vast quantities of information--far more than the visual system can process in a short period of time-and spatial attention is therefore used to focus the visual system's processing resources onto a subset of the incoming visual information. Most psychological theories of attention posit a single mechanism for this focusing of attention, but recent electrophysiological studies have provided evidence that the visual system employs several separable neural mechanisms of spatial attention. This paper describes the evidence for multiple attentional mechanisms and suggests links between these neurophysiologically defined mechanisms and specific functional processes that have been proposed in psychological theories of attention.     

Attentional bias to brief threat-related faces revealed by saccadic eye movements.

According to theories of emotion and attention, we are predisposed to orient rapidly toward threat. However, previous examination of attentional cueing by threat showed no enhanced capture at brief durations, a finding that may be related to the sensitivity of the manual response measure used. Here we investigated the time course of orienting attention toward fearful faces in the exogenous cueing task. Cue duration (20 ms or 100 ms) and response mode (saccadic or manual) were manipulated. In the saccade mode, both enhanced attentional capture and impaired disengagement from fearful faces were evident and limited to 20 ms, suggesting that saccadic cueing effects emerge rapidly and are short lived. In the manual mode, fearful faces impacted only upon the disengagement component of attention at 100 ms, suggesting that manual cueing effects emerge over longer periods of time. Importantly, saccades could reveal threat biases at brief cue durations consistent with current theories of emotion and attention. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Challenging the anterior attentional system with a continuous performance task: a functional magnetic resonance imaging approach

Combining the Continuous Performance Test (CPT) with a modern functional imaging technique provides a powerful tool for investigating neurophysiological processes in the human brain. There is increasing evidence from single photon emission tomography (SPECT), positron emission tomography (PET) and presently also functional magnetic resonance imaging (fMRI) studies proposing the existence of a distributed large-scale attentional network, mediated by the dorsolateral prefrontal and mesial frontal cortex, thalamus, basal ganglia and posterior parietal and superior temporal lobe. The aim of this study is to show that fMRI is a useful tool for in vivo localization of attentional tasks and to compare the results with established imaging techniques. Functional MRI was performed on a clinical 1.5-T system using gradient-echo acquisition. For data processing, the Statistical Parametric Mapping (SPM96) package was used. A right lateralized activation pattern in the dorsolateral prefrontal and mesial frontal cortex, the thalamus and the basal ganglia was found in a group of 12 male subjects. These findings support theories suggesting right hemispheric dominance of human attention.