Anling Rao

Synergistic effect of combined temporal and spatial expectations on visual attention.

We developed a naturalistic behavioral task to investigate the influence of spatial and temporal expectations on attentional orienting to moving targets. In this task, the movement of an object before its disappearance under an occluding barrier generated expectations concerning the location and/or time of its reappearance. Four different trial types were intermixed, each inducing a different state of expectation: no expectation, only spatial expectation about the location of reappearance, only temporal expectation about the moment of reappearance, and combined spatial and temporal expectation. The behavioral validity of the task was shown by the fact that all expectation conditions produced significantly shorter reaction times than the control state of no expectation. Spatial attention modulated early perceptual analysis in extrastriate areas, as demonstrated by significant enhancement of the visual P1 component. Temporal attention alone had no effect on P1 but instead modulated response-specific components. However, when spatial and temporal attention were combined, the enhancement of perceptual processing was significantly augmented, leading to a greater enhancement of the P1 component than by spatial attention alone. Perceptual analysis reflected by the P1 component correlated significantly with reaction times. In summary, event-related potentials revealed the presence of individual modulatory effects attributable to spatial and temporal expectation as well as synergistic effects indicative of an interaction of the two. This synergistic effect is likely to play a critical role in directing attention to the reappearance of a temporarily occluded moving target, a process of obvious importance in everyday situations.

Searching for Targets within the Spatial Layout of Visual Short-Term Memory

Recent studies have revealed that the internal representations that we construct from the environment and maintain in visual short-term memory (VSTM) to guide behavior are highly flexible and can be selectively modulated according to our task goals and expectations. In the current study, we conducted two experiments to compare and contrast neural mechanisms of selective attention related to searching for target items within perceptual versus VSTM representations. We used event-related potentials to investigate whether searching for relevant target items from within VSTM representations involves spatially specific biasing of neural activity in a manner analogous to that which occurs during visual search for target items in perceptual arrays. The results, replicated across the two experiments, revealed that selection of a target object within a search array maintained in VSTM proceeds through a similar mechanism as that in the perceptual domain. In line with previous results, N2pc potentials were obtained when targets were identified within a perceptual visual-search array. Interestingly, equivalent N2pcs, with similar time courses and scalp distributions, were also elicited when target items were identified within a VSTM representation. The findings reinforce the notion of highly flexible VSTM representations that can be modulated according to task goals and suggest a large degree of overlap in the spatially specific neural mechanisms of target selection across the perceptual and VSTM domains.

Spatial attention can bias search in visual short-term memory.

Whereas top-down attentional control is known to bias perceptual functions at many levels of stimulus analysis, its possible influence over memory-related functions remains uncharted. Our experiment combined behavioral measures and event-related potentials (ERPs) to test the ability of spatial orienting to bias functions associated with visual short-term memory (VSTM), and to shed light on the neural mechanisms involved. In particular, we investigated whether orienting attention to a spatial location within an array maintained in VSTM could facilitate the search for a specific remembered item. Participants viewed arrays of one, two or four differently colored items, followed by an informative spatial (100% valid) or uninformative neutral retro-cue (1500–2500 ms after the array), and later by a probe stimulus (500–1000 ms after the retro-cue). The task was to decide whether the probe stimulus had been present in the array. Behavioral results showed that spatial retro-cues improved both accuracy and response times for making decisions about the presence of the probe item in VSTM, and significantly attenuated performance decrements caused by increasing VSTM load. We also identified a novel ERP component (N3RS) specifically associated with searching for an item within VSTM. Paralleling the behavioral results, the amplitude and duration of the N3RS systematically increased with VSTM load in neutral retro-cue trials. When spatial retro-cues were provided, this “retro-search” component was absent. Our findings clearly show that the influence of top-down attentional biases extends to mnemonic functions, and, specifically, that searching for items within VSTM can be under flexible voluntary control.

Selective Attention to Specific Features within Objects: Behavioral and Electrophysiological Evidence

Evidence regarding the ability of attention to bias neural processing at the level of single features has been gathering steadily, but most of the experiments to date used arrays with multiple objects and locations, making it difficult to rule out indirect influences from object or spatial attention. To investigate feature-specific selective attention, we have assessed the ability to select and ignore individual features within the same object. We used a negative-priming paradigm in which the color or the direction of internal motion of the object could determine the relevant response. Bidimensional (colored and moving) and unidimensional (colored and stationary, or gray and moving) stimuli appeared in unpredictable order. In successive blocks, participants were instructed that one feature dimension was dominant. During that block, participants responded according to the dominant dimension for bidimensional stimuli. For unidimensional stimuli, participants responded to the only dimension of the stimulus that afforded a response, regardless of the instruction for the block. The ability to inhibit irrelevant task information at the level of specific features (negative priming for features) was indexed by a decrease in performance to detect one particular feature value (e.g., red) if the same feature value (red) but not another color value (green) had been ignored in the previous bidimensional stimulus. Behavioral results confirmed the existence of inhibitory, negative-priming mechanisms at the singlefeature level for both color and motion dimensions of stimuli. Event-related potentials recorded during task performance revealed the dynamics of neural modulation by feature attention. Comparisons were made using the identical physical stimuli under different conditions of attention to isolate purely attentional effects. Processing of identical bidimensional stimuli was compared as a function of the dimension of attention (color, motion). Processing of identical unidimensional stimuli that followed bidimensional stimuli was also compared to identify possible effects of feature-specific negative priming. The electrophysiological effects revealed that inhibition of irrelevant features leads to modulation of brain activity during early stages of perceptual analysis.

Anticipating conflict facilitates controlled stimulus-response selection

Cognitive control can be triggered in reaction to previous conflict, as suggested by the finding of sequential effects in conflict tasks. Can control also be triggered proactively by presenting cues predicting conflict (“proactive control”)? We exploited the high temporal resolution of ERPs and controlled for sequential effects to ask whether proactive control based on anticipating conflict modulates neural activity related to cognitive control, as may be predicted from the conflict-monitoring model. ERPs associated with conflict detection (N2) were measured during a cued flanker task. Symbolic cues were either informative or neutral with respect to whether the target involved conflicting or congruent responses. Sequential effects were controlled by analyzing the congruency of the previous trial. The results showed that cueing conflict facilitated conflict resolution and reduced the N2 latency. Other potentials (frontal N1 and P3) were also modulated by cueing conflict. Cueing effects were most evident after congruent than after incongruent trials. This interaction between cueing and sequential effects suggests neural overlap between the control networks triggered by proactive and reactive signals. This finding clarifies why previous neuroimaging studies, in which reactive sequential effects were not controlled, have rarely found anticipatory effects upon conflict-related activity. Finally, the high temporal resolution of ERPs was critical to reveal a temporal modulation of conflict detection by proactive control. This novel finding suggests that anticipating conflict speeds up conflict detection and resolution. Recent research suggests that this anticipatory mechanism may be mediated by preactivation of ACC during the preparatory interval.

The effects of combined caffeine and glucose drinks on attention in the human brain.

Abstract The objective of this research was to measure the effects of energising drinks containing caffeine and glucose, upon mental activity during sustained selective attention. Non-invasive electrophysiological brain recordings were made during a behavioural study of selective attention in which participants received either energising or placebo drinks. We tested specifically whether energising drinks have significant effects upon behavioural measures of performance during a task requiring sustained visual selective attention, as well as on accompanying components of the event-related potential (ERPs) related to information processing in the brain. Forty healthy volunteers were blindly assigned to receive either the energising drink or a similar-tasting placebo drink. The behavioural task involved identifying predefined target stimulus among rapidly presented streams of peripheral visual stimuli, and making speeded motor responses to this stimulus. During task performance, accuracy, reaction times and ongoing brain activity were stored for analysis. The energising drink enhanced behavioural performance both in terms of accuracy and speed of reactions. The energising drink also had significant effects upon the event-related potentials. Effects started from the enhancement of the earliest components (C1/P1), reflecting early visual cortical processing in the energising-drink group relative to the placebo group over the contralateral scalp. The later N1, N2 and P3 components related to decision-making and responses were also modulated by the energising drink. Energising drinks containing caffeine and glucose can enhance behavioural performance during demanding tasks requiring selective attention. The behavioural benefits are coupled to direct effects upon neural information processing.

Watching where you look: Modulation of visual processing of foveal stimuli by spatial attention

Two experiments investigated the effect of sustained selective spatial attention upon the perceptual analysis of stimuli within the center of gaze. Spatial attention has typically been studied in relation to peripheral stimuli, and its relevance to the processing of central stimuli has remained relatively ignored. Here, behavioral measures in normal human volunteer participants showed that focused spatial attention can also influence responses to central stimuli, over and beyond the advantage conferred by their foveation. Event-related potentials (ERPs) showed that the action of attention upon foveal stimuli included the modulation of perceptual processing in extrastriate visual areas. Surprisingly, the visual modulation revealed an intriguing and consistent pattern of hemispheric asymmetry, in both experiments. These findings show that in addition to the established right hemisphere dominance of the brain areas that direct attention, the consequences of directed attention may also be asymmetrical.

Biasing Perception by Spatial Long-Term Memory

Human perception is highly flexible and adaptive. Selective processing is tuned dynamically according to current task goals and expectations to optimize behavior. Arguably, the major source of our expectations about events yet to unfold is our past experience; however, the ability of long-term memories to bias early perceptual analysis has remained untested. We used a noninvasive method with high temporal resolution to record neural activity while human participants detected visual targets that appeared at remembered versus novel locations within naturalistic visual scenes. Upon viewing a familiar scene, spatial memories changed oscillatory brain activity in anticipation of the target location. Memory also enhanced neural activity during early stages of visual analysis of the target and improved behavioral performance. Both measures correlated with subsequent target-detection performance. We therefore demonstrated that memory can directly enhance perceptual functions in the human brain.

Modulation of neural activity by motivational and spatial biases.

Motivational biases and spatial attention both modulate neural activity and influence behavioural performance. The time course of motivational bias effects, as well as the relationship between motivation and attention across the time course of information processing, however, are relatively unknown. In the present study, event-related potentials (ERPs) were recorded whilst individuals performed a modified Posner task, in which cue stimuli indicated the reward stakes of a given trial and the probable spatial location of a subsequent target stimulus. Reaction times (RTs) were sensitive to motivation and to attention, with faster responses produced on valid and on rewarded trials. In addition, motivation modulated neural activity from the visual analysis of stimuli, with an earlier N1 peak for rewarded compared with non-rewarded stimuli. Effects of motivation were relatively independent from those of attention until late cognitive processing and response production, where motivation and attention interacted to enhance P300-like potentials and the lateralised readiness potential (LRP). The results suggest that multiple sources of modulatory influences may exist, with motivation and attention exerting independent influences over early stimulus and cognitive processing, followed by a late interaction allowing the construction of a comprehensive stimulus representation that contains information pertaining to both motivational and spatial expectations.

Brain potentials associated with conscious aftereffects induced by unseen stimuli in a blindsight subject

The study is of brain activity in a blindsight subject (D.B.), who reports conscious visual afterimages of stimuli of which he is unaware when they are presented. This contrast offered a unique opportunity to study event-related potential recordings of conscious versus unconscious visual phenomena generated by the very same stimulus in the identical locus of the visual field. The behavioral results confirmed the reliability of the difference in the subjects report for inducing stimuli versus their aftereffects. The rationale of the event-related potential analysis was to subtract “on” signals from “off” signals, the latter associated with the onset of conscious events and the former for events that remained unconscious. Because there are inherent differences in on and off potentials, the subtractive resultants for the blind hemifield were compared with the same subtractions for the good hemifield when the subject was aware both of the stimuli and their afterimages. A differential pattern in subtractive resultants emerged with a strong anterior left frontal focus for the blind field and a posterior focus for the intact field. The results are compared with other studies suggesting an anterior focus for conscious visual events.