Dissociable forms of inhibitory control within prefrontal cortex with an analog of the Wisconsin Card Sort Test: restriction to novel situations and independence from "on-line" processing

Attentional set-shifting and discrimination reversal are sensitive to prefrontal damage in the marmoset in a manner qualitatively similar to that seen in man and Old World monkeys, respectively (Dias et al., 1996b). Preliminary findings have demonstrated that although lateral but not orbital prefrontal cortex is the critical locus in shifting an attentional set between perceptual dimensions, orbital but not lateral prefrontal cortex is the critical locus in reversing a stimulus-reward association within a particular perceptual dimension (Dias et al., 1996a). The present study presents this analysis in full and extends the results in three main ways by demonstrating that (1) mechanisms of inhibitory control and "on-line" processing are independent within the prefrontal cortex, (2) impairments in inhibitory control induced by prefrontal damage are restricted to novel situations, and (3) those prefrontal areas involved in the suppression of previously established response sets are not involved in the acquisition of such response sets. These findings suggest that inhibitory control is a general process that operates across functionally distinct regions within the prefrontal cortex. Although damage to lateral prefrontal cortex causes a loss of inhibitory control in attentional selection, damage to orbitofrontal cortex causes a loss of inhibitory control in affective processing. These findings provide an explanation for the apparent discrepancy between human and nonhuman primate studies in which disinhibition as measured on the Wisconsin Card Sort Test is associated with dorsolateral prefrontal damage, whereas disinhibition as measured on discrimination reversal is associated with orbitofrontal damage.     

The role of spatial and nonspatial information in visual selection.

Even though it is undisputed that prior information regarding the location of a target affects visual selection, the issue of whether information regarding nonspatial features, such as color and shape, has similar effects has been a matter of debate since the early 1980s. In the study described in this article, measures derived from signal detection theory were used to show that perceptual sensitivity is affected by a top-down set for spatial information but not by a top-down set for nonspatial information. This indicates that knowing where the target singleton is affects perceptual selectivity but that knowing what it is does not help selectivity. Furthermore, perceptual sensitivity can be enhanced by nonspatial features, but only through a process related to bottom-up priming. These findings have important implications for models of visual selection. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Spatial, but not object, delayed response is impaired in early Parkinson's disease.

The authors hypothesized that the pathophysiology of early Parkinson's disease (PD) may selectively target structures that support visual working memory for spatial relations but leave structures that support working memory for featural characteristics of objects relatively intact. Fifteen PD and 15 normal control participants took a visual delayed-response test with a spatial condition and a (nonspatial) object condition, equating the perceptual difficulty of the tests for each participant. The stimuli were irregular polygons presented at different locations on a computer screen. Results revealed a selective impairment of spatial delayed response in PD, indicating a disruption of spatial working memory unconfounded by sensory processing difficulties. The selectivity of this deficit may reflect the circumscribed nature of pathophysiological change affecting the caudate nucleus in early PD. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Intradimensional and extradimensional shifts in spatial learning.

Animals trained on 2 discriminations learn the 2nd rapidly if the relevant stimuli are from the same dimension as the 1st (an intradimensional or ID shift) but slowly if the relevant stimuli for the 2 problems are from different dimensions (an extradimensional or ED shift). Four experiments examined ID and ED shifts in spatial learning. Rats trained on 2 spatial problems learned the 2nd more rapidly than rats whose 1st problem had been nonspatial. But this difference between ID and ED shifts depended on the spatial relationship between rewarded (S+) and unrewarded (S-) alternatives in the 2 spatial problems. The results imply that rats trained on a spatial discrimination do not learn to attend to all spatial landmarks but only to those that serve to differentiate S+ and S-. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The influence of temporal selection on spatial selection and distractor interference: An attentional blink study.

Both spatial and temporal selection require focused attention. The authors examine how temporal attention affects spatial selection. In a dual-task rapid serial visual presentation paradigm, temporal selection of a target (T1) impairs processing of a second target (T2) that follows T1 within 500 ms. This process is the attentional blink (AB). To test the effects of withdrawing temporal attention, the authors measured concurrent distractor interference on T2 when the distractors were presented during and outside of the AB. Perceptual interference was manipulated by the similarity in color between T2 and concurrent distractors, and response interference was manipulated by the flanker congruency task. Results showed that perceptual interference was larger during the AB. Response interference also increased during the AB, but only when perceptual interference was high. The authors conclude that temporal selection and spatial selection rely on a common attentional process. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The limit of tactile spatial resolution in humans: grating orientation discrimination at the lip, tongue, and finger

The sensory neural pathways serving the lip, tongue, and finger are specialized for spatial information processing; thus, damage to these pathways is likely to be manifested most prominently as a loss of spatial acuity. For that reason, accurate measurement of spatial resolution at these regions is particularly important. The conventional test, the two-point discrimination task, does not measure the limit of spatial resolution and it yields variable results because it does not control nonspatial cues. The aim of this study was to quantify the limits of spatial resolution at the lip, tongue, and finger and to study the repeatability of those measurements using a stimulus that does not introduce nonspatial cues. We employed a grating orientation discrimination test, which has been studied extensively in relation to the underlying neural mechanisms. We obtained psychophysical thresholds for tactile spatial resolution from 15 normal, young adult subjects over seven test sessions. The finest gratings whose orientations were discriminated reliably had groove widths (gratings had equal groove and bar widths) that averaged 0.51 mm at the lip, 0.58 mm at the tongue, and 0.94 mm at the finger. These threshold measurements were highly reproducible between sessions with an overall improvement of 2% per session. These data suggest that the grating orientation discrimination task provides a stable, reliable measure of the human capacity for spatial resolution.     

Why does a red snake in the grass capture your attention?

It has been suggested that affective stimuli automatically capture attention; this preferential processing is thought to be related to the evolutionary significance of affective stimuli. However, recent evidence suggests that perceptual salience alone might explain why some affective stimuli are more likely to influence attentional processes in certain contexts. In this study, we manipulated affective and perceptual salience to better understand how affective information is processed and how it impacts attentional processes in different contexts. We used stimuli that are both affectively and perceptually salient, while varying the task requirement to encourage the processing of perceptually salient (Experiment 1) or affectively salient (Experiment 2) information. This design made it possible to observe independent and interdependent relationships between perceptual and affective salience. The results showed that when the task encouraged the processing of perceptually salient information, affective salience did not influence task performance. In contrast, when the task encouraged the processing of affectively salient information, affectively salient information impaired task performance. The findings suggest that the affective nature of the stimuli does not always influence attentional processes. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

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)     

The influence of alertness on spatial and nonspatial components of visual attention.

Three experiments investigated whether spatial and nonspatial components of visual attention would be influenced by changes in (healthy, young) subjects’ level of alertness and whether such effects on separable components would occur independently of each other. The experiments used a no-cue/alerting-cue design with varying cue-target stimulus onset asynchronies in two different whole-report paradigms based on Bundesen’s (1990) theory of visual attention, which permits spatial and nonspatial components of selective attention to be assessed independently. The results revealed the level of alertness to affect both the spatial distribution of attentional weighting and processing speed, but not visual short-term memory capacity, with the effect on processing speed preceding that on the spatial distribution of attentional weighting. This pattern indicates that the level of alertness influences both spatial and nonspatial component mechanisms of visual attention and that these two effects develop independently of each other; moreover, it suggests that intrinsic and phasic alertness effects involve the same processing route, on which spatial and nonspatial mechanisms are mediated by independent processing systems that are activated, due to increased alertness, in temporal succession. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Implicit affective cues and attentional tuning: An integrative review.

A large and growing number of studies support the notion that arousing positive emotional states expand, and that arousing negative states constrict, the scope of attention on both the perceptual and conceptual levels. However, these studies have predominantly involved the manipulation or measurement of conscious emotional experiences (e.g., subjective feelings of happiness or anxiety). This raises the question: Do cues that are merely associated with benign versus threatening situations but do not elicit conscious feelings of positive or negative emotional arousal independently expand or contract attentional scope? Integrating theoretical advances in affective neuroscience, positive psychology, and social cognition, the authors propose that rudimentary intero- and exteroceptive stimuli may indeed become associated with the onset of arousing positive or negative emotional states and/or with appraisals that the environment is benign or threatening and thereby come to moderate the scope of attention in the absence of conscious emotional experience. Specifically, implicit “benign situation” cues are posited to broaden, and implicit “threatening situation” cues to narrow, the range of both perceptual and conceptual attentional selection. An extensive array of research findings involving a diverse set of such implicit affective cues (e.g., enactment of approach and avoidance behaviors, incidental exposure to colors signaling safety vs. danger) is marshaled in support of this proposition. Potential alternative explanations for and moderators of these attentional tuning effects, as well as their higher level neuropsychological underpinnings, are also discussed along with prospective extensions to a range of other situational cues and domains of social cognitive processing. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Electrophysiological markers of visual dimension changes and response changes.

In cross-dimensional visual search tasks, target discrimination is faster when the previous trial contained a target defined in the same visual dimension as the current trial. The dimension-weighting account (DWA; A. Found & H. J. Müller, 1996) explains this intertrial facilitation by assuming that visual dimensions are weighted at an early perceptual stage of processing. Recently, this view has been challenged by models claiming that intertrial facilitation effects are generated at later stages that follow attentional target selection (K. Mortier, J. Theeuwes, & P. A. Starreveld, 2005). To determine whether intertrial facilitation is generated at a perceptual stage, at the response selection stage, or both, the authors focused on specific event-related brain potential components (directly linkable to perceptual and response-related processing) during a compound search task. Visual dimension repetitions were mirrored by shorter latencies and enhanced amplitudes of the N2-posterior- contralateral, suggesting a facilitated allocation of attentional resources to the target. Response repetitions and changes systematically modulated the lateralized readiness potential amplitude, suggesting a benefit from residual activations of the previous trial biasing the correct response. Overall, the present findings strengthen the DWA by indicating a perceptual origin of dimension change costs in visual search. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Discrimination learning and attentional set formation in a mouse model of Fragile X.

Fragile X Syndrome is the most prevalent genetic cause of mental retardation. Selective deficits in executive function, including inhibitory control and attention, are core features of the disorder. In humans, Fragile X results from a trinucleotide repeat in the Fmr1 gene that renders it functionally silent and has been modeled in mice by targeted deletion of the Fmr1 gene. Fmr1 knockout (KO) mice recapitulate many features of Fragile X syndrome, but evidence for deficits in executive function is inconsistent. To address this issue, we trained wild-type and Fmr1 KO mice on an experimental paradigm that assesses attentional set-shifting. Mice learned to discriminate between stimuli differing in two of three perceptual dimensions. Successful discrimination required attending only to the relevant dimension, while ignoring irrelevant dimensions. Mice were trained on three discriminations in the same perceptual dimension, each followed by a reversal. This procedure normally results in the formation of an attentional set to the relevant dimension. Mice were then required to shift attention and discriminate based on a previously irrelevant perceptual dimension. Wild-type mice exhibited the increase in trials to criterion expected when shifting attention from one perceptual dimension to another. In contrast, the Fmr1 KO group failed to show the expected increase, suggesting impairment in forming an attentional set. Fmr1 KO mice also exhibited a general impairment in learning discriminations and reversals. This is the first demonstration that Fmr1 KO mice show a deficit in attentional set formation. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

Is there feature-based attentional selection in visual search?

A new paradigm combines attentional cuing and rapid serial visual presentation to disentangle the effects of perceptual filtering and location selection. Observers search successive, superimposed arrays, in which feature values are alternated for a target numeral among letters. Two dimensions, size (small, large) and color (red, green) are tested. Selective attention to feature values is jointly manipulated by instructions, presentation probabilities, and payoffs. In Experiment 1, the attended feature provides temporal, not spatial, information; observers show no attentional costs or benefits in response accuracy. In Experiment 2, the attended feature indicates a unique location; observers show consistent attentional costs and benefits. Selective attention to a particular size or color does not cause perceptual exclusion or admission of items containing that feature; it acts by guiding search processes to spatial locations that contain the to-be-attended feature. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Primate analogue of the Wisconsin card sorting test: Effects of excitotoxic lesions of the prefrontal cortex in the marmoset.

Using a primate analogue of the Wisconsin Card Sort Test, this study demonstrated, for the first time, that lesions of the prefrontal cortex in monkeys produce a qualitatively similar impairment in attentional set-shifting to that seen following prefrontal cortical damage in humans. Although damage to the prefrontal cortex did not disrupt the ability of marmosets, a New World monkey, to maintain an attentional set, it did disrupt their ability to shift an attentional set. It also impaired their performance on discrimination reversal, object retrieval, and spatial delayed response, consistent with the effects of prefrontal damage in Old World monkeys. Comparison of the cognitive processes underlying discrimination reversal, object retrieval, and attentional set-shifting reveals the various types of inhibitory control provided by the prefrontal cortex. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Object exploration and reactions to spatial and nonspatial changes in hooded rats following damage to parietal cortex or hippocampal formation.

Hooded rats with bilateral lesions of the anterior part of the hippocampal formation (HIP), anterior region of the posterior parietal cortex (APC), or posterior region of the posterior parietal cortex (PPC) were compared with controls for their exploration of 5 objects in an open field, habituation of locomotion and object investigation, and response to spatial and nonspatial change. First, all groups displayed habituation of both locomotor and exploratory activity. Second, controls selectively reexplored displaced objects, and APC-lesioned rats reexplored all objects, whereas PPC- and HIP-lesioned rats failed to react to the spatial change. Third, a novel object induced reexploration in all groups. Results are consistent with the roles of the HIP and PPC in spatial information processing. Moreover, the APC and PPC are involved in attentional effortful processing and visuospatial information processing necessary for spatial representation, respectively. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Covert auditory spatial orienting: An evaluation of the spatial relevance hypothesis.

The spatial relevance hypothesis (J. J. McDonald & L. M. Ward, 1999) proposes that covert auditory spatial orienting can only be beneficial to auditory processing when task stimuli are encoded spatially. We present a series of experiments that evaluate 2 key aspects of the hypothesis: (a) that “reflexive activation of location-sensitive neurons is not sufficient to produce attentional facilitation” and (b) that “any task constraint that makes space important for the listener will produce auditory spatial cue effects” (p. 1236). Experiment 1 showed significant reflexive-orienting benefits on a nonspatial task, refuting the first claim. However, Experiments 2 to 4 reveal that informative spatial cues can improve performance on a nonspatial task, consistent with the second claim. Auditory spatial-cue benefits found with nonspatial tasks appear smaller and less reliable than those found in visual spatial-orienting studies, possibly due to differences in the coding of spatial information in vision and audition. The final experiments consider the mechanisms by which auditory spatial orienting might facilitate auditory processing and provide tentative evidence that attention enhances processing at one ear rather than influencing neurons tuned to the attended location. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

When visual marking meets the attentional blink: More evidence for top-down, limited-capacity inhibition.

An attentional blink (AB) paradigm was used to investigate the attentional resources necessary for visual marking. The results showed that distractors presented inside the AB cannot easily be ignored despite participants anticipating a future target display. This supports the hypothesis that attentional resources are required for visual marking. In addition, probe dots were better detected on blinked distractors than on successfully ignored distractors, but only when the task required new items to be prioritized. In a final experiment, a stronger negative carry-over effect on search occurred for targets identical to distractors presented outside rather than inside the AB. This suggests that at least part of the inhibitory processes involved in visual marking are nonspatial. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Resolving Visual Interference During Covert Spatial Orienting: Online Attentional Control Through Static Records of Prior Visual Experience.

Models of attentional control usually describe online shifts in control settings that accommodate changing task demands. The current studies suggest that online control over distractor exclusion--a core component of visual selection--can be accomplished without online shifts in top-down settings. Measurements of target discrimination accuracy suggested that the degree of distractor exclusion was guided by retinotopic maps of the prior probability of distractor interference at the attended locations. These probability maps can be retrieved via object-based cues, and they interact with shifts of attention to elicit increased levels of distractor exclusion when it is most needed. Thus, static probability maps can provide an internal template that guides the resolution of visual interference as spatial attention traverses the visual field. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The structure of attentional control: Contingent attentional capture by apparent motion, abrupt onset, and color.

Five spatial cuing experiments tested 2 hypotheses regarding attentional capture: (1) Attentional capture is contingent on endogenous attentional control settings, and (2) attentional control settings are limited to the distinction between dynamic and static discontinuities as proposed by C. L. Folk et al (see record 1993-12300-001). In Exps 1 and 2, apparent-motion precues produced significant costs in performance for targets signaled by motion but not for targets signaled by color or abrupt onset. Exp 3 established that this pattern is not due to differences in the difficulty of target discrimination. Exps 4 and 5 revealed asymmetric capture effects between abrupt onset and apparent motion related to stimulus salience. Results support the hypotheses of Folk et al and suggest that stimulus salience may also play a role in attentional capture. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Attentional switching in spatial and nonspatial domains: Evidence from the attentional blink.

Much is known abut attritional switching across space, but much less about switches between nonspatial domains such as category or task. Nonetheless, extensive information about attentional switching in both spatial and nonspatial domains can be found in the experimental literature on a phenomenon known as the attentional blink, in which a switch is required between 2 rapidly sequential targets. If the two targets follow one another directly, identification of the second target is almost perfect when no attentional switch is required between the targets or when the switch is unidimensional. In contrast, identification is impaired with switches in location or with multidimensional switches. This pattern of results is consistent with the joint operation of location-specific endogenously controlled input filters and exogenously controlled domain-specific modules. (PsycINFO Database Record (c) 2010 APA, all rights reserved)