Effects of altering brain cholinergic activity on covert orienting of attention: comparison of monkey and human performance

Experiments were conducted to elucidate the role of the cholinergic neurotransmitter system in arousal and the orienting of attention to peripheral targets. Rhesus monkeys and humans fixated a visual stimulus and responded to the onset of visual targets presented randomly in two visual field locations. The target was preceded by a valid cue (cue and target at the same location), an invalid cue (cue and target to opposite locations), a double cue (cues to both spatial locations, target to one), or, the cue was omitted (no-cue, target to either location). Reaction times (RTs) to the onset of the target were recorded. For monkeys, systemic injections of nicotine (0.003-0.012 mg/kg) or atropine (0.001-0.01 mg/kg), but not saline control injections, reduced mean RTs for all trials, indicating general behavioral stimulation. In addition, nicotine significantly reduced RTs for invalid trials but had little additional effect on those for valid, double, or no-cue trials. Virtually identical effects were observed for human chronic tobacco smokers in performing the same task following cigarette smoking. Injections of atropine in monkeys had no effect on RTs for valid or invalid trials but significantly slowed RTs in double-cue trials that did not require the orienting of attention. These results suggest that in both species, the nicotinic cholinergic system may play a role in automatic sensory orienting. In addition, the muscarinic system may play a role in alerting to visual stimuli in monkeys.     

Inhibition of return along the path of attention.

Investigated inhibition of return (IOR) at 4 target locations (a near and a far location in each visual hemifield). Exp 1 was conducted to determine if IOR extends over a cued visual hemifield. 12 Ss indicated when they visually detected a cross after being instructed to keep their eyes fixed on a specific point. The target locations were aligned horizontally and IOR was observed at the near locations when cues were presented at the far locations but not at the far locations when cues were presented at the near locations. In Exp 2, the target locations were not horizontally aligned and IOR was not observed for near locations when the far locations were cued. Otherwise, the same procedure and apparatus as in Exp 1 were used by 12 different Ss. The authors conclude that IOR may be an attentional phenomenon and that attention may operate in an analog fashion such that locations in the path of the attentional movement may be attended to. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Sustained visual-spatial attention produces costs and benefits in response time and evoked neural activity

This study investigated the simple reaction time (RT) and event-related potential (ERP) correlates of biasing attention towards a location in the visual field. RTs and ERPs were recorded to stimuli flashed randomly and with equal probability to the left and right visual hemifields in the three blocked, covert attention conditions: (i) attention divided equally to left and right hemifield locations; (ii) attention biased towards the left location; or (iii) attention biased towards the right location. Attention was biased towards left or right by instructions to the subjects, and responses were required to all stimuli. Relative to the divided attention condition, RTs were significantly faster for targets occurring where more attention was allocated (benefits), and slower to targets where less attention was allocated (costs). The early P1 (100-140 msec) component over the lateral occipital scalp regions showed attentional benefits. There were no amplitude modulations of the occipital N1 (125-180 msec) component with attention. Between 200 and 500 msec latency, a late positive deflection (LPD) showed both attentional costs and benefits. The behavioral findings show that when sufficiently induced to bias attention, human observers demonstrate RT benefits as well as costs. The corresponding P1 benefits suggest that the RT benefits of spatial attention may arise as the result of modulations of visual information processing in the extrastriate visual cortex.     

The retinotopy of visual spatial attention

We used high-field (3T) functional magnetic resonance imaging (fMRI) to label cortical activity due to visual spatial attention, relative to flattened cortical maps of the retinotopy and visual areas from the same human subjects. In the main task, the visual stimulus remained constant, but covert visual spatial attention was varied in both location and load. In each of the extrastriate retinotopic areas, we found MR increases at the representations of the attended target. Similar but smaller increases were found in V1. Decreased MR levels were found in the same cortical locations when attention was directed at retinotopically different locations. In and surrounding area MT+, MR increases were lateralized but not otherwise retinotopic. At the representation of eccentricities central to that of the attended targets, prominent MR decreases occurred during spatial attention.     

Is Posner's "beam" the same as Treisman's "glue?" On the relation between visual orienting and feature integration theory.

In the present study we investigated whether the visually allocated "beam" studied by Posner and others is the same visual attentional resource that performs the role of feature integration in Treisman's model. Subjects were cued to attend to a certain spatial location by a visual cue, and performance at expected and unexpected stimulus locations was compared. Subjects searched for a target letter (R) with distractor letters that either could give rise to illusory conjunctions (PQ) or could not (PB). Results from three separate experiments showed that orienting attention in response to central cues (endogenous orienting) showed similar effects for both conjunction and feature search. However, when attention was oriented with peripheral visual cues (exogenous orienting), conjunction search showed larger effects of attention than did feature search. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Shifts of visual attention to multiple simultaneous location cues.

Conducted 2 experiments to examine the effects of adjacent double cueing on target identification response times (Exp 1) and to determine whether similar results would be obtained with nonadjacent double cueing (Exp 2). For 14 observers, single and simultaneous multiple location cues were presented a brief period of time before the onset of targets that they were required to identify. Single and multiple cueing effects were similar, even when targets were presented at locations "in between" pairs of cues. The increase in response times that occurred when a target was presented at uncued locations in between a pair of cued locations suggests that pairs of cued locations are encoded independently by an operation called spatial indexing. A multiple indexing proposal and its role in mediating visual attention shifts initiated by direct cueing is described. (French abstract) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Bilateral interactions in saccade programming. A saccade-latency study

Subjects were required to make a saccade to a target appearing randomly 4 degree to the left or right of the current fixation position (1280 trials per experiment). Location cues were used to direct visual attention and start saccade preparation to one of the two locations before target onset. When the cue indicated the target location (valid trials), the generation of express saccades (visually guided saccades with latencies around 100 ms) was strongly facilitated. When the opposite location was cued (invalid trials), express saccades were abolished and replaced by a population of mainly fast-regular saccades (latencies around 150 ms). This was found with a peripheral cue independently of whether the fixation point was removed before target onset (gap condition; experiment 1) or remained on throughout the trial (overlap condition; experiment 2). The same pattern also was observed with a central cue that did not involve any visual stimulation at a peripheral location (experiment 3). In the case where the primary saccade was executed in response to the cue and the target appeared at the opposite location, continuous amplitude transition functions were observed: starting at about 60-70 ms from target onset onward, the amplitude of the cue-elicited saccades continuously decreased from 4 degree to values below 1 degree. The results are explained by a fixation-gating model, according to which the antagonism between fixation and saccade activity gives rise to multimodal distributions of saccade latencies. It is argued that allocation of visual attention and saccade preparation to one location entails a successive disengagement of the fixation system controlling saccade preparation within the hemifield to which the saccade is prepared and a partial engagement of the opposite fixation system.     

Spatiotemporal distribution of facilitation and inhibition of return arising from the reflexive orienting of covert attention.

Currently, there is debate regarding both the spatial and temporal relationship between facilitation and inhibition of return (IOR) components of attention, as observed on the covert orienting of visual attention task (COVAT). These issues were addressed in a series of experiments where the spatial and temporal relationships between cue and target were manipulated. Facilitation occurred only when the stimulus onset asynchrony (SOA) was short and there was temporal overlap between cue and target. IOR occurred only when SOA was long and there was no temporal overlap between cue and target. Facilitation encompassed the cued location and all locations between the cue and fixation, whereas IOR arose for the entire cued hemifield. These findings suggest that the facilitation and IOR found on COVATs that use noninformative peripheral cues are separable and stimulus-driven processes. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Perceptually based implicit learning in severe closed-head injury patients.

This study suggests that perceptually based implicit learning may either be preserved following a severe closed-head injury (CHI) or recover within the 1st year. Nineteen severe CHI patients and 19 controls searched visual matrices and indicated the quadrant location of a target. Participants were exposed to the following covariation pattern: AAAABAAA. For Covariation A blocks, the matrices systematically co-occurred with a unique location of the target. This relationship was altered for the B block. Despite CHI participants' overall slower response times (RTs), both groups demonstrated the expected decline in RTs across the first 4 Covariation A blocks followed by an increase when the covariation changed. Both groups also exhibited retention of their learning after a 20-min delay. Explicit knowledge tests indicated that participants lacked awareness for the covariation. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Spatial cuing in a stereoscopic display: attention remains "depth-aware" with age

Previous research has demonstrated that spatial attention is "depth-aware": Reaction times (RT) are greater for shifts in depth and two-dimensional (2-D) space than in 2-D space alone. This experiment examined whether the ability to focus attention at a depth location is maintained with advanced age. Twelve younger and 12 older observers viewed stereoscopic displays in which one of four spatial locations was cued. Two of the locations were at a near depth location and two were at a far depth location. When the focus of visual attention was shifted to a new location in space (because of an invalid cue), the cost in RT for switching attention (measured as the difference between RT on valid cue and invalid cue trials) was greater when observers had to switch attention between different depth locations and different locations in 2-D space than for shifts in 2-D space alone. This effect was observed for both younger and older observers, suggesting that the ability to orient attention to a depth location is maintained with age.     

Target location probability effects in visual search: An effect of sequential dependencies.

Target location probability was manipulated in a visual search task. When the target was twice as likely to appear on 1 side of the display as the other, manual button-press response times were faster (Experiment 1A) and first saccades were more frequently directed (Experiment 1B) to the more probable locations. When the target appeared with equal probability at each location in this search task, performance benefited from repetition of target location in the preceding trials (Experiment 2). When the trial sequence was constrained so that target location did not repeat within a series of 4 trials, there was no longer an advantage for more probable locations (Experiment 3). The authors conclude that the search benefits for more probable locations resulted from short-term target location repetitions. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The role of the fixation location in inhibition of return.

In examining the role of the fixation location in inhibition of return (IOR), 3 experiments were conducted in which an exogenous cue was used to reorient attention following a peripheral cue and before the appearance of a target. However, this cue occurred at either the traditional central fixation location or a nonfixated location. In Exp 1, 10 undergraduates' eye positions were monitored at the 3 different times during the trial (after the onset of the cue at 1 of the target locations; after the onset of the 2nd cue; after the onset of the target). In Exp 2, 16 undergraduates repeated the same procedures as in Exp 1 except that an additional cue was added. In Exp 3, 8 undergraduates repeated the same procedures as in Exp 1 except the placement of the fixation, peripheral, and target locations was altered. The results indicate that reorienting attention to a fixated location results in a significant reduction in the inhibitory effect. The results from the study suggest that IOR could serve as a mechanism that improves the efficiency of visual searches. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Attentional priming and visual search in pigeons.

Advance information about a target's identity improved visual search efficiency in pigeons. Experiments 1 and 2 compared information supplied by visual cues with information supplied by trial sequences. Reaction times (RTs) were lower when visual cues signaled a single target rather than two. RTs were lower (Experiment 1) or accuracy improved (Experiment 2) when a sequence of trials presented a single target rather than a mixture of 2. Experiments 3, 4, and 5 considered the selectivity of visual priming by introducing probe trials that reversed the usual cue–target relationship. RT was higher following such miscues than following the usual 1- or 2-target cuing relationships (Experiment 3); the miscuing effect persisted over variations in the target's concealment (Experiments 4 and 5), but did not occur when the target was presented alone (Experiment 4). Findings indicate that priming modifies an attentional mechanism and suggest that this effect accounts for search images. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Consciousness of attention and expectancy as reflected in event-related potentials and reaction times.

The effects of conscious expectancies and attention on event-related potentials (ERPs) and choice reaction times (RTs) and their modulation by stimulus sequence were studied. Ss retrospectively reported their expectancy of, and attention to, the terminal tones of short series. ERPs and RTs showed the usual sequential effects that were modulated by practice. As ratings were affected by only a few of the stimulus sequence, conscious access to sequence-based expectancy or attention appears to be fragmentary. Increased P300 amplitude with attention indicates conscious access to processing capacity. RTs and P300 latencies suggest stimulus processing time to decrease with sequence-based and consciously accessible expectancy. Differential effects of stimulus sequences and conscious expectancies on P300 amplitude indicate influences of two varieties of expectancy. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Auditory and audiovisual inhibition of return

Two experiments examined any inhibition-of-return (IOR) effects from auditory cues and from preceding auditory targets upon reaction times (RTs) for detecting subsequent auditory targets. Auditory RT was delayed if the preceding auditory cue was on the same side as the target, but was unaffected by the location of the auditory target from the preceding trial, suggesting that response inhibition for the cue may have produced its effects. By contrast, visual detection RT was inhibited by the ipsilateral presentation of a visual target on the preceding trial. In a third experiment, targets could be unpredictably auditory or visual, and no peripheral cues intervened. Both auditory and visual detection RTs were now delayed following an ipsilateral versus contralateral target in either modality on the preceding trial, even when eye position was monitored to ensure central fixation throughout. These data suggest that auditory target-target IOR arises only when target modality is unpredictable. They also provide the first unequivocal evidence for cross-modal IOR, since, unlike other recent studies (e.g., Reuter-Lorenz, Jha, & Rosenquist, 1996; Tassinari & Berlucchi, 1995; Tassinari & Campara, 1996), the present cross-modal effects cannot be explained in terms of response inhibition for the cue. The results are discussed in relation to neurophysiological studies and audiovisual links in saccade programming.     

Combined expectancies: Event-related potentials reveal the early benefits of spatial attention that are obscured by reaction time measures.

Visual spatial attention has been likened to a "spotlight" that selectively facilitates the perceptual processing of events at covertly attended locations. However, if participants have advance knowledge of the likely location of an impending target and the likely response it will require, facilitation in response performance does not occur for targets at the expected (or attended) location that require an unexpected response. Event-related potentials (ERPs) were recorded during a discrimination task in which the most likely target location and target response were simultaneously cued prior to target onset. The ERPs showed evidence of enhanced perceptual-level processing for all targets at attended locations. These results suggest that the lack of response facilitation for unexpected targets at attended locations is likely due to postperceptual processes that are activated by the inclusion of nonspatial stimulus expectancies, response expectancies, or both. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Asymmetric object substitution masking.

Object substitution masking occurs when a lateral mask persists beyond the duration of a target, reflecting reentrant processes in vision (V. Di Lollo, J. Enns, & R. Rensink, 2000). The authors studied whether substitution masking is location specific and whether it is symmetric around the target. A brief circular display of letters was presented along with a mask that designated the target. The mask was centered on the target or 1.1° to the central or to the peripheral side. Substitution masking was found even when the target and the mask were at different locations. It was asymmetric and stronger when the mask was to the peripheral side of the target than to the central side. Asymmetric substitution was observed using various masks. It could not be explained by retina acuity gradients and was not attenuated by focused attention. The authors propose that target selection triggers an asymmetric inhibitory surround that is stronger toward the central side of the target. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Development of orienting to locations and objects in human infants.

Investigated the preference for orienting to novel locations and novel objects in young infants in 2 experiments, by examining the influence of visual signals on subsequent attentional orienting and eye movements. Exp 1 (n?=?12) demonstrates that 3-mo-olds show inhibition of return (IOR) for 10° target eccentricities, but not for 30° target eccentricities. In Exp 2, 14 3-mo-olds and 14 6-mo-olds oriented to 10° targets that varied in location and object identity. Ss of both ages strongly preferred orienting to novel objects at novel locations. At 3 mo, the preference for novel objects was equal to the preference for novel locations, while at 6 mo a tendency to prefer novel objects over novel locations emerged. Overall, findings support separate development of these 2 forms of novelty preference, and suggest that novel location preferences (IOR) relates closely to the eye movement system. (French abstract) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Gating of neuronal responses in macaque primary visual cortex by an attentional spotlight

Neuronal responses were recorded from the striate cortex of monkeys trained to perform visual discrimination at locations in the visual field to which their attention was drawn. A subset of neurons showed vigorous responses to visual stimuli for trials in which the monkey was directing its attention to the respective receptive field location. In trials where attention is directed elsewhere, responses to the same stimuli were significantly reduced. In some cells the early response component was not modulated by attention, but later components were affected by the locus of attention. The results suggest the operation of a feedback in the paradigm that spotlights a topographically restricted area of V1 for further processing at higher levels.     

Saccade preparation inhibits reorienting to recently attended locations.

We measured manual reaction time in normal human Ss to confirm that an eccentric visual signal has a biphasic effect on covert attention and eye movements. First, it summons attention and biases a saccade toward the signal; a subsequent inhibition of return then slows responses to signals at that location. A temporal hemifield dominance for inhibition of return was shown; this finding coverges with observations (M. I. Posner et al; see record 1986-22316-001) in neurologic patients to suggest that it is mediated by midbrain pathways. Endogenous orienting of attention, from a central arrow cue, did not activate inhibition of return, whereas endogenous saccade preparation did so as effectively as an exogenous signal, even when no saccade was made. Inhibition of return is activated by midbrain oculomotor pathways and may function as a location "tagging" mechanism to optimize efficiency of visual search. (PsycINFO Database Record (c) 2010 APA, all rights reserved)