The orienting of visuospatial attention: an event-related brain potential study

This study investigated the electrophysiological correlates of shifting, maintaining, and relaxing the focus of attention, using a symbolic cuing task. Cues and imperative stimuli were presented in rapid succession, and the ADJAR procedure was used to remove the contribution of event-related potential (ERP) activity associated with the imperative stimulus from the cue-related ERP waveforms. Initial analyses, comparing left and right attention-directing cues, replicated previous findings of early directing attention negativity (EDAN) and anterior directing attention negativity (ADAN) effects. To isolate ERP activity that is common to leftward and rightward attention shifts, the combined ERP activity elicited by attention-directing cues was compared to the ERP activity elicited by non-informative cues. This analysis revealed a strong and broadly distributed early positivity followed by a sustained central negativity, possibly reflecting the controlled orienting and subsequent maintenance of attentional focus. Finally, imperative stimuli preceded by non-informative cues were characterized by an enhanced posterior P2 effect, with a scalp distribution indicative of generators in visual areas. This result suggests a relatively late (re)activation in visual areas associated with the processing of stimuli that had not been cued in advance.     

Recoding between two types of STM representation revealed by the dynamics of memory search

Visual STM (VSTM) is thought to be related to visual attention in several ways. Attention controls access to VSTM during memory encoding and plays a role in the maintenance of stored information by strengthening memorized content. We investigated the involvement of visual attention in recall from VSTM. In two experiments, we measured electrophysiological markers of attention in a memory search task with varying intervals between VSTM encoding and recall, and so we were able to track recoding of representations in memory. Results confirmed the involvement of attention in VSTM recall. However, the amplitude of the N2pc and N3rs components, which mark orienting of attention and search within VSTM, decreased as a function of delay. Conversely, the amplitude of the P3 and sustained posterior contralateral negativity components increased as a function of delay, effectively the opposite of the N2pc and N3rs modulations. These effects were only observed when verbal memory was not taxed. Thus, the results suggested that gradual recoding from visuospatial orienting of attention into verbal recall mechanisms takes place from short to long retention intervals. Interestingly, recall at longer delays was faster than at short delays, indicating that verbal representation is coupled with faster responses. These results extend the orienting-of-attention hypothesis by including an account of representational recoding during short-term consolidation and its consequences for recall from VSTM.     

Attention modulates maintenance of representations in visual short-term memory

Recent studies have shown that selective attention is of considerable importance for encoding task-relevant items into visual short-term memory (VSTM) according to our behavioral goals. However, it is not known whether top-down attentional biases can continue to operate during the maintenance period of VSTM. We used ERPs to investigate this question across two experiments. Specifically, we tested whether orienting attention to a given spatial location within a VSTM representation resulted in modulation of the contralateral delay activity (CDA), a lateralized ERP marker of VSTM maintenance generated when participants selectively encode memory items from one hemifield. In both experiments, retrospective cues during the maintenance period could predict a specific item (spatial retrocue) or multiple items (neutral retrocue) that would be probed at the end of the memory delay. Our results revealed that VSTM performance is significantly improved by orienting attention to the location of a task-relevant item. The behavioral benefit was accompanied by modulation of neural activity involved in VSTM maintenance. Spatial retrocues reduced the magnitude of the CDA, consistent with a reduction in memory load. Our results provide direct evidence that top-down control modulates neural activity associated with maintenance in VSTM, biasing competition in favor of the task-relevant information.     

Neural activity associated with attention orienting triggered by gaze cues: A study of lateralized ERPs

This study investigated the electrophysiological (event-related potential, ERP) and behavioral (reaction time, RT) correlates of gaze-cued shifts of visuospatial attention. Participants viewed centrally presented faces with neutral expressions in which eyes looked straight ahead and then shifted to the left or right. Upon movement of the eyes, the facial expression either stayed the same (neutral) or changed to a fearful or happy expression. Participants' task was to identify a probe letter (T or L), which appeared in either the same or the opposite location to the direction of the eye gaze. There was behavioral evidence of a gaze congruency effect, as RTs were faster when the eyes looked toward rather than away from the location of the target. The ERP data indicated the presence of a significant gaze-congruent anterior directing attention negativity (ADAN) at anterior locations (300–500 ms after the onset of the gaze cue). ERP data did not show evidence of initial orienting of attention triggered by gaze cues in the early directing attention negativity (EDAN) at posterior locations (200–300 ms post-cue onset). The gaze cueing effects in the RT and ERP data were not significantly influenced by the emotional expression of the faces. The presence of the ADAN may reflect neural mechanisms that underlie the holding of attention on gazed-at locations.     

An ERP study of preparatory and inhibitory mechanisms in a cued saccade task

The present study uses event-related potentials (ERPs) to investigate the neurophysiological correlates of the mechanisms involved in selection of locations for saccades. Participants performed a task in which both target and distractor locations were cued on a trial-by-trial basis. Participants were instructed to make an eye movement to the cued target location and to ignore elements at the distractor location. This experimental set-up allowed the investigation of the mechanisms involved in the top-down preparation and inhibition of locations for an eye movement. When comparing responses to leftward and rightward pointing cues, we observed an early directing attention negativity (EDAN) and an anterior directing attention negativity (ADAN) effect in the cue-target interval. These effects were similar to those observed in studies investigating attentional allocation, suggesting a close link between shifts of spatial attention and the preparation of eye movements. These components were followed by a late widespread contralateral negativity (LDAN) that was assumed to reflect both the oculomotor programming of the upcoming eye movement as well as attentional orienting. Furthermore, a new component was observed related to top-down inhibition of the distractor location. In response to the distractor cue, an early positivity above the right hemisphere (RLIP) was revealed. Finally, no modulations of early target-evoked ERP components were observed, suggesting that these components are unaffected when no further processing is required at the cued location.     

Effects of attentional filtering demands on preparatory ERPs elicited in a spatial cueing task

ObjectiveWe used ERP measures to investigate how attentional filtering requirements affect preparatory attentional control and spatially selective visual processing.MethodsIn a spatial cueing experiment, attentional filtering demands were manipulated by presenting task-relevant visual stimuli either in isolation (target-only task) or together with irrelevant adjacent distractors (target-plus-distractors task). ERPs were recorded in response to informative spatial precues, and in response to subsequent visual stimuli at attended and unattended locations.ResultsThe preparatory ADAN component elicited during the cue-target interval was larger and more sustained in the target-plus-distractors task, reflecting the demand of stronger attentional filtering. By contrast, two other preparatory lateralised components (EDAN and LDAP) were unaffected by the attentional filtering demand. Similar enhancements of P1 and N1 components in response to the lateral imperative visual stimuli were observed at cued versus uncued locations, regardless of filtering demand, whereas later attentional-related negativities beyond 200 ms post-stimulus were larger the target-plus-distractor task.ConclusionsOur results implicate that the ADAN component is linked to preparatory top-down control processes involved in the attentional filtering of irrelevant distractors; such filtering also affects later attention-related negativities recorded after the onset of the imperative stimulus.SignificanceERPs can reveal effects of expected attentional filtering of irrelevant distractors on preparatory attentional control processes and spatially selective visual processing.     

Effects of hand posture on preparatory control processes and sensory modulations in tactile-spatial attention.

OBJECTIVE: Event-related brain potentials (ERPs) were measured to investigate spatial coordinate systems involved in the control of preparatory tactile-spatial orienting, and in subsequent attentional modulations of somatosensory processing. METHODS: On each trial, a visual precue directed attention to the left or right hand, where infrequent tactile targets had to be detected. Hands were positioned either close together or wide apart. ERPs were recorded in the cue-target interval and in response to attended and unattended tactile non-targets. RESULTS: A frontal anterior directing attention negativity (ADAN) and a posterior late directing attention positivity (LDAP) were elicited in the cue-target interval contralateral to the direction of an attentional shift. The ADAN was unaffected by hand posture, but the LDAP was attenuated when hands were close together. N140 amplitudes were enhanced in response to tactile stimuli presented to the attended hand, and this effect was more pronounced when hands were wide apart. CONCLUSIONS: ADAN and LDAP are linked to separable anterior and posterior attentional control systems, which use coordinate systems based on somatotopic and external space, respectively. Effects of spatial attention on somatosensory stimulus processing are affected by variations in body posture. SIGNIFICANCE: Our results demonstrate that representations of body locations in external space play a central role in the control of tactile attention.     

Top-down visual activity underlying VSTM and preparatory attention

Attention and visual short-term memory (VSTM) are both associated with top-down activity in visual cortex. To-be-remembered visual input triggers persistent activity in visual cortex, and preparatory attention elicits visual activity in anticipation of an expected target stimulus. This brief review considers similarities, and differences, in top-down visual activity underlying VSTM maintenance and preparatory attention. The extant evidence suggests that top-down visual activity provides a common neural substrate for VSTM maintenance and preparatory attention, however the precise neural implementation depends on the specific task parameters. A similar neurobiological framework may also apply to other high-level visual phenomena, including visual imagery and awareness.     

On the equivalence of executed and imagined movements: Evidence from lateralized motor and nonmotor potentials

The neural simulation theory assumes that motor imagery and motor execution draw on a shared set of mechanisms underlying motor cognition. Evidence is accumulating that motor imagery and motor execution have many common features. The extent of the similarity and whether it spreads into the preparation phase is however unclear. This study used electroencephalographic recordings to compare the effects of providing advance information about upcoming movements on preparatory processing in a motor imagery and execution paradigm. Event‐related potential data were recorded in a priming task where participants were cued to perform simple or complex finger movements. We hypothesized that a high degree of functional similarity of motor imagery and motor execution should be reflected in similar alterations of lateralized preparatory activity. Lateralized preparatory activity was indeed very similar, showing both motor‐related (lateralized readiness potential, LRP) and cognitive components (anterior directing‐attention negativity or ADAN, late directing‐attention positivity or LDAP). Dipole analysis revealed that LRP, ADAN, and LDAP sources were very comparable for motor imagination and execution. Results generally support the idea of common underlying functional networks subserving both the preparation for execution and imagery of movements. They also provide a broader context for this notion by revealing similarities in cognitive components associated with the movement tasks. Hum Brain Mapp, 2009. © 2009 Wiley‐Liss, Inc.     

Selective attention and the three-process memory model for the interpretation of verbal free recall in amyotrophic lateral sclerosis

The present study investigates selective attention and verbal free recall in amyotrophic lateral sclerosis (ALS) and examines the contribution of selective attention, encoding, consolidation, and retrieval memory processes to patients' verbal free recall. We examined 22 non-demented patients with sporadic ALS and 22 demographically related controls using Stroop Neuropsychological Screening Test (SNST; selective attention) and Rey Auditory Verbal Learning Test (RAVLT; immediate & delayed verbal free recall). The item-specific deficit approach (ISDA) was applied to RAVLT to evaluate encoding, consolidation, and retrieval difficulties. ALS patients performed worse than controls on SNST (p < .001) and RAVLT immediate and delayed recall (p < .001) and showed deficient encoding (p = .001) and consolidation (p = .002) but not retrieval (p = .405). Hierarchical regression analysis revealed that SNST and ISDA indices accounted for: (a) 91.1% of the variance in RAVLT immediate recall, with encoding (p = .016), consolidation (p < .001), and retrieval (p = .032) significantly contributing to the overall model and the SNST alone accounting for 41.6%; and (b) 85.2% of the variance in RAVLT delayed recall, with consolidation (p < .001) and retrieval (p = .008) significantly contributing to the overall model and the SNST alone accounting for 39.8%. Thus, selective attention, encoding, and consolidation, and to a lesser extent of retrieval, influenced both immediate and delayed verbal free recall. Concluding, selective attention and the memory processes of encoding, consolidation, and retrieval should be considered while interpreting patients' impaired free recall. (JINS, 2012, 18, 1-10).     

Encoding: the keystone to efficient functioning of verbal short-term memory

Verbal short-term memory (VSTM) is thought to play a critical role in language learning. It is indexed by the nonword repetition task where listeners are asked to repeat meaningless words like ‘blonterstaping’. The present study investigated the effect on nonword repetition performance of differences in efficiency of functioning of some part of the neural architecture mediating VSTM. Hypotheses were stated within Baddeley and Hitch's (1974) multicomponent model of VSTM, with respect to regions of the brain known to be active during tasks tapping into VSTM. We were specifically interested in activations associated with the posterior planum temporale (Spt) which emerge during rehearsal since this region is hypothesized to be central to VTSM (Buchsbaum, Olsen, Koch, & Berman, 2005a).Participants performed a delayed reaction time task in the scanner which explicitly mimicked the three main stages of information-processing involved in VSTM (encoding, rehearsal, recall (here recognition)). The data for each stage were then convolved with scores from a separately measured nonword repetition task.Rather than observing a pattern of individual differences located to specific regions specialized for supporting VSTM, a dissociation in direction of correlation in overlapping regions of the brain was observed during encoding and recognition. Larger hemodynamic responses during encoding were associated with better nonword repetition, and vice versa during recognition. There was little evidence for a network of activations specialized for VSTM. Instead, the main correlations were observed in regions also known to be involved in long-term memory. It seems that individuals who are better at nonword repetition and hence at language learning, activate these regions more efficiently than poorer nonword-repeaters early after stimulus input. These observations are discussed with respect to various models proposed for explaining the phenomenon of VSTM.     

Clinical and cognitive factors associated with verbal memory task performance in patients with schizophrenia

OBJECTIVE: The authors have previously shown the role of depression, slowing of processing speed, and selective attention deficit in verbal memory task performance in schizophrenia. They wished to determine the specific contribution of each of these factors to various types of memory impairment. METHOD: The negative symptom score from the Positive and Negative Syndrome Scale, the Hamilton Depression Rating Scale score, a measure of processing speed, and a measure of selective attention were entered as predictors in regression analyses. Furthermore, analyses of covariance were conducted on the memory measures to test the significance of the differences between schizophrenic patients and healthy comparison subjects after control for processing speed and selective attention. RESULTS: Depression was associated only with deep encoding reflected by semantic clustering. Selective attention was associated only with superficial encoding reflected by serial recall. Slowing of processing speed was associated with both deep and superficial encoding. Negative symptoms were not associated with memory impairment except for the avolition item from the Scale for the Assessment of Negative Symptoms. Processing speed accounted for all the group differences on the memory measures that reflected superficial encoding. In addition, a subgroup of patients with no or minor depression was not significantly impaired on deep encoding relative to the healthy comparison group. CONCLUSIONS: The authors suggest that verbal memory impairment in schizophrenia is a consequence of depression and slowness, rather than a primary feature of the disease.     

The role of biased competition in visual short-term memory

Despite being extensively studied, many important questions regarding visual short-term memory (VSTM) are yet to be fully answered. The present review seeks to explore the extent to which competitive interactions present during perception of stimuli may also operate during stimulus encoding and/or maintenance in VSTM. Additionally, the paper provides an overview of the methods and approaches that have been used to study the properties of VSTM, and a review of research into the limits of VSTM capacity. We take as our starting point the biased competition model of attention (Desimone & Duncan, 1995) and discuss the influence that such competition may have on the limited capacity of VSTM by a review of the literature and by recent experiments from Shapiro's lab. The present report outlines several experiments that provide results consistent with the idea that low-level competitive interactions may influence VSTM, with the aim of stimulating further research into this new area.     

Manipulations of attention dissociate fragile visual short-term memory from visual working memory

People often rely on information that is no longer in view, but maintained in visual short-term memory (VSTM). Traditionally, VSTM is thought to operate on either a short time-scale with high capacity - iconic memory - or a long time scale with small capacity - visual working memory. Recent research suggests that in addition, an intermediate stage of memory in between iconic memory and visual working memory exists. This intermediate stage has a large capacity and a lifetime of several seconds, but is easily overwritten by new stimulation. We therefore termed it fragile VSTM. In previous studies, fragile VSTM has been dissociated from iconic memory by the characteristics of the memory trace. In the present study, we dissociated fragile VSTM from visual working memory by showing a differentiation in their dependency on attention. A decrease in attention during presentation of the stimulus array greatly reduced the capacity of visual working memory, while this had only a small effect on the capacity of fragile VSTM. We conclude that fragile VSTM is a separate memory store from visual working memory. Thus, a tripartite division of VSTM appears to be in place, comprising iconic memory, fragile VSTM and visual working memory.     

Distinct electrophysiological indices of maintenance in auditory and visual short-term memory

We compared the electrophysiological correlates for the maintenance of non-musical tones sequences in auditory short-term memory (ASTM) to those for the short-term maintenance of sequences of coloured disks held in visual short-term memory (VSTM). The visual stimuli yielded a sustained posterior contralateral negativity (SPCN), suggesting that the maintenance of sequences of coloured stimuli engaged structures similar to those involved in the maintenance of simultaneous visual displays. On the other hand, maintenance of acoustic sequences produced a sustained negativity at fronto-central sites. This component is named the Sustained Anterior Negativity (SAN). The amplitude of the SAN increased with increasing load in ASTM and predicted individual differences in the performance. There was no SAN in a control condition with the same auditory stimuli but no memory task, nor one associated with visual memory. These results suggest that the SAN is an index of brain activity related to the maintenance of representations in ASTM that is distinct from the maintenance of representations in VSTM.     

Spatiotemporal overlap between brain activation related to saccade preparation and attentional orienting

Recent brain imaging studies provided evidence that the brain areas involved with attentional orienting and the preparation of saccades largely overlap, which may indicate that focusing attention at a specific location can be considered as an unexecuted saccade towards that location (i.e. the premotor theory of attention). Alternatively, it may be proposed that attentional orienting is simply relevant for preparing saccades, but the two processes may also be completely unrelated. In two experiments, we examined temporal activation of brain areas by measuring the electroencephalogram. Central cues indicated the likely side (left or right) at which a to-be-attended target would occur, or to which a saccade had to be prepared. Cue direction-related activity was determined, time-locked to cue onset. In addition, in our second experiment, delayed saccades had to be carried out, which allows to focus on processes strongly related to saccade execution. In nearly all tasks, an early directing attention negativity (EDAN), an anterior directing attention negativity (ADAN), and a late directing attention positivity (LDAP) were observed, time-locked to cue onset. Source analyses supported the view that this activity probably originates from areas within the ventral intraparietal sulcus (vIPS) and the frontal eye fields (FEF). The saccade-locked analysis also indicated that the FEF plays an important role in triggering saccades, but the role of vIPS appears to be minimal. The latter finding disfavors the premotor theory of attention, as it suggests that the relation between attention and action is less direct.     

Remembered but unused: the accessory items in working memory that do not guide attention

If we search for an item, a representation of this item in our working memory guides attention to matching items in the visual scene. We can hold multiple items in working memory. Do all these items guide attention in parallel? We asked participants to detect a target object in a stream of objects while they maintained a second item in memory for a subsequent task. On some trials, we presented this memory item as a distractor in the stream. Subjects did not confuse these memory items with the search target, as the false alarm rate on trials where the memory item was presented in the stream was comparable to that on trials with only regular distractors. However, a comparable performance does not exclude that the memory items are processed differently from normal distractors. We therefore recorded event-related potentials (ERPs) evoked by search targets, memory items, and regular distractors. As expected, ERPs evoked by search targets differed from those evoked by distractors. Search targets elicited an occipital selection negativity and a frontal selection positivity indexing selective attention, whereas the P3b component, which reflects the matching of sensory events to memory representations, was enhanced for targets compared to distractors. Remarkably, the ERPs evoked by memory items were indistinguishable from the ERPs evoked by normal distractors. This implies that the search target has a special status in working memory that is not shared by the other items. These other, "accessory" items do not guide attention and are excluded from the matching process.     

Rapid targeting followed by sustained deployment of visual spatial attention

We investigated preparatory attention processes when a spatial discrimination was required at a cued location, by measuring electroencephalography following a central symbolic cue to deploy spatial attention. Electroencephalography activity in response to the cue revealed three cue-related activations: an early-onset positivity following the P1 at posterior scalp sites contralateral to the cued location, followed by cue-related frontal scalp activity and later-onset sustained activity at posterior scalp sites contralateral to the cued location. The early contralateral positivity may reflect rapid targeting of the cued location. Our results also extend the findings of cue-related frontal activity followed by posterior activity contralateral to the cued location, found with nonspatial feature discriminations, to a task requiring a spatial discrimination.     

Shifts of attention in the early blind: an erp study of attentional control processes in the absence of visual spatial information

To investigate the role of visual spatial information in the control of spatial attention, event-related brain potentials (ERPs) were recorded during a tactile attention task for a group of totally blind participants who were either congenitally blind or had lost vision during infancy, and for an age-matched, sighted control group who performed the task in the dark. Participants had to shift attention to the left or right hand (as indicated by an auditory cue presented at the start of each trial) in order to detect infrequent tactile targets delivered to this hand. Effects of tactile attention on the processing of tactile events, as reflected by attentional modulations of somatosensory ERPs to tactile stimuli, were very similar for early blind and sighted participants, suggesting that the capacity to selectively process tactile information from one hand versus the other does not differ systematically between the blind and the sighted. ERPs measured during the cue-target interval revealed an anterior directing attention negativity (ADAN) that was present for the early blind group as well as for the sighted control group. In contrast, the subsequent posterior late direction attention negativity (LDAP) was absent in both groups. These results suggest that these two components reflect functionally distinct attentional control mechanisms which differ in their dependence on the availability of visually coded representations of external space.     

Memory and schizophrenia: differential link of processing speed and selective attention with two levels of encoding

The purpose of this study was to investigate how underlying cognitive deficits such as a defect in processing speed or in selective attention contributed to different types of memory impairment observed in schizophrenia (superficial vs deep encoding). 49 schizophrenic patients and 40 normal controls were administered a verbal memory task. Superficial encoding was assessed by the ability to recall items in their serial order. Deep encoding was assessed by the ability to organise words into semantic categories. Two measures of processing speed (Digit Symbol Substitution Test and Stroop colour time) and one measure of selective attention (Stroop test) were used. Regression analyses were carried out. In the patient group, processing speed contributed to both superficial and deep encoding, and to a global verbal memory score. Selective attention only contributed to the superficial encoding processes. Thus, slowing of processing speed in schizophrenia seems to be more crucial for memory performance, since it affects memory in a pervasive way.