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Todd C. Handy
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Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2014) 26 (3): 635–644.
Published: 01 March 2014
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Our ability to make decisions is predicated upon our knowledge of the outcomes of the actions available to us. Reinforcement learning theory posits that actions followed by a reward or punishment acquire value through the computation of prediction errors—discrepancies between the predicted and the actual reward. A multitude of neuroimaging studies have demonstrated that rewards and punishments evoke neural responses that appear to reflect reinforcement learning prediction errors [e.g., Krigolson, O. E., Pierce, L. J., Holroyd, C. B., & Tanaka, J. W. Learning to become an expert: Reinforcement learning and the acquisition of perceptual expertise. Journal of Cognitive Neuroscience, 21, 1833–1840, 2009; Bayer, H. M., & Glimcher, P. W. Midbrain dopamine neurons encode a quantitative reward prediction error signal. Neuron, 47, 129–141, 2005; O'Doherty, J. P. Reward representations and reward-related learning in the human brain: Insights from neuroimaging. Current Opinion in Neurobiology, 14, 769–776, 2004; Holroyd, C. B., & Coles, M. G. H. The neural basis of human error processing: Reinforcement learning, dopamine, and the error-related negativity. Psychological Review, 109, 679–709, 2002]. Here, we used the brain ERP technique to demonstrate that not only do rewards elicit a neural response akin to a prediction error but also that this signal rapidly diminished and propagated to the time of choice presentation with learning. Specifically, in a simple, learnable gambling task, we show that novel rewards elicited a feedback error-related negativity that rapidly decreased in amplitude with learning. Furthermore, we demonstrate the existence of a reward positivity at choice presentation, a previously unreported ERP component that has a similar timing and topography as the feedback error-related negativity that increased in amplitude with learning. The pattern of results we observed mirrored the output of a computational model that we implemented to compute reward prediction errors and the changes in amplitude of these prediction errors at the time of choice presentation and reward delivery. Our results provide further support that the computations that underlie human learning and decision-making follow reinforcement learning principles.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2013) 25 (6): 952–960.
Published: 01 June 2013
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Mind wandering is a natural, transient state wherein our neurocognitive systems become temporarily decoupled from the external sensory environment as our thoughts drift away from the current task at hand. Yet despite the ubiquity of mind wandering in everyday human life, we rarely seem impaired in our ability to adaptively respond to the external environment when mind wandering. This suggests that despite widespread neurocognitive decoupling during mind wandering states, we may nevertheless retain some capacity to attentionally monitor external events. But what specific capacities? In Experiment 1, using traditional performance measures, we found that both volitional and automatic forms of visual–spatial attentional orienting were significantly attenuated when mind wandering. In Experiment 2, however, ERPs revealed that, during mind wandering states, there was a relative preservation of sensitivity to deviant or unexpected sensory events, as measured via the auditory N1 component. Taken together, our findings suggest that, although some selective attentional processes may be subject to down-regulation during mind wandering, we may adaptively compensate for these neurocognitively decoupled states by maintaining automatic deviance–detection functions.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2011) 23 (12): 3725–3733.
Published: 01 December 2011
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Previous research has demonstrated that higher-order cognitive processes associated with the allocation of selective attention are engaged when highly familiar self-relevant items are encountered, such as one's name, face, personal possessions and the like. The goal of our study was to determine whether these effects on attentional processing are triggered on-line at the moment self-relevance is established. In a pair of experiments, we recorded ERPs as participants viewed common objects (e.g., apple, socks, and ketchup) in the context of an “ownership” paradigm, where the presentation of each object was followed by a cue indicating whether the object nominally belonged either to the participant (a “self” cue) or the experimenter (an “other” cue). In Experiment 1, we found that “self” ownership cues were associated with increased attentional processing, as measured via the P300 component. In Experiment 2, we replicated this effect while demonstrating that at a visual–perceptual level, spatial attention became more narrowly focused on objects owned by self, as measured via the lateral occipital P1 ERP component. Taken together, our findings indicate that self-relevant attention effects are triggered by the act of taking ownership of objects associated with both perceptual and postperceptual processing in cortex.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2011) 23 (2): 460–470.
Published: 01 February 2011
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Top–down control of visual sensory cortex has long been tied to the orienting of visual spatial attention on a rapid, moment-to-moment basis. Here, we examined whether sensory responses in visual cortex are also modulated by natural and comparatively slower fluctuations in whether or not one is paying attention to the task at hand. Participants performed a simple visual discrimination task at fixation as the ERPs to task-irrelevant probes in the upper visual periphery were recorded. At random intervals, participants were stopped and asked to report on their attentional state at the time of stoppage—either “on-task” or “off-task.” ERPs to the probes immediately preceding these subjective reports were then examined as a function of whether attention was in an on-task versus off-task state. We found that sensory-evoked responses to the probes were significantly attenuated during off-task relative to on-task states, as measured by the visual P1 ERP component. In two additional experiments, we replicated this effect while (1) finding that off-task sensory attenuation extends to the auditory domain, as measured by the auditory N1 ERP component, and (2) eliminating state-dependent shifts in general arousal as a possible explanation for the effects. Collectively, our findings suggest that sensory gain control in cortex is yoked to the natural ebb and flow in how much attention we pay to the current task over time.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2010) 22 (1): 124–138.
Published: 01 January 2010
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We know that human neurocognitive systems rapidly and implicitly evaluate emotionally charged stimuli. But what about more everyday, frequently encountered kinds of objects, such as computer desktop icons and business logos? Do we rapidly and implicitly evaluate these more prosaic visual images, attitude objects that might only engender a mild sense of liking or disliking, if at all? To address this question, we asked participants to view a set of unfamiliar commercial logos in the context of a target identification task as brain electrical responses to these objects were recorded via event-related potentials (ERPs). Following this task, participants individually identified those logos that were most liked or disliked, allowing us to then compare how ERP responses to logos varied as a function of hedonic evaluation—a procedure decoupling evaluative responses from any normative classification of the logos themselves. In Experiment 1, we found that visuocortical processing manifest a specific bias for disliked logos that emerged within the first 200 msec of stimulus onset. In Experiment 2, we replicated this effect while dissociating normative- and novelty-related influences. Taken together, our results provide direct electrophysiological evidence suggesting that we rapidly and implicitly evaluate commercial branding images at a hedonic level.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2008) 20 (5): 879–891.
Published: 01 May 2008
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This study examines whether orienting attention to biologically based social cues engages neural mechanisms distinct from those engaged by orienting to nonbiologically based nonsocial cues. Participants viewed a perceptually ambiguous stimulus presented centrally while performing a target detection task. By having participants alternate between viewing this stimulus as an eye in profile or an arrowhead, we were able to directly compare the neural mechanisms of attentional orienting to social and nonsocial cues while holding the physical stimulus constant. The functional magnetic resonance imaging results indicated that attentional orienting to both eye gaze and arrow cues engaged extensive dorsal and ventral fronto-parietal networks. Eye gaze cues, however, more vigorously engaged two regions in the ventral frontal cortex associated with attentional reorienting to salient or meaningful stimuli, as well as lateral occipital regions. An event-related potential study demonstrated that this enhanced occipital response was attributable to a higher-amplitude sensory gain effect for targets appearing at locations cued by eye gaze than for those cued by an arrowhead. These results endorse the hypothesis that differences in attention to social and nonsocial cues are quantitative rather than qualitative, running counter to current models that assume enhanced processing for social stimuli reflects the involvement of a unique network of brain regions. An intriguing implication of the present study is the possibility that our ability to orient volitionally and reflexively to socially irrelevant stimuli, including arrowheads, may have arisen as a useful by-product of a system that developed first, and foremost, to promote social orienting to stimuli that are biologically relevant.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2008) 20 (3): 458–469.
Published: 01 March 2008
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Converging evidence from neuroscience suggests that our attention to the outside world waxes and wanes over time. We examined whether these periods of “mind wandering” are associated with reduced cortical analysis of the external environment. Participants performed a sustained attention to response task in which they responded to frequent “nontargets” (digits 0–9) and withheld responses for infrequent “targets” (the letter X). Mind wandering was defined both behaviorally, indicated by a failure to withhold a response to a target, and subjectively, via self-report at a thought probe. The P300 event-related potential component for nontargets was reduced prior to both the behavioral and subjective reports of mind wandering, relative to periods of being “on-task.” Regression analysis of P300 amplitude revealed significant common variance between behavioral and subjective markers of mind wandering, suggesting that both markers reflect a common underlying mental state. Finally, control analysis revealed that the effect of mind wandering on the P300 could not be ascribed to changes in motor activity nor was it associated with general arousal. Our data suggest that when trying to engage attention in a sustained manner, the mind will naturally ebb and flow in the depth of cognitive analysis it applies to events in the external environment.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2005) 17 (12): 1936–1949.
Published: 01 December 2005
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Attention-related sensory gain control in human extra-striate cortex is believed to improve the acuity of visual perception. Yet given wide variance in the spatial resolution of vision across the retina, it remains unclear whether sensory gain operates homogenously between foveal and nonfoveal retinotopic locations. To address this issue, we used event-related potentials (ERPs) in a variant of the canonical spatial attention task. Participants were cued to expect targets at either fixation (foveal targets) or at a location several degrees above fixation (parafoveal targets). At both target locations, manual reaction times were shorter for cued relative to uncued targets, indicating that attention was consistently oriented to the cued location. Nevertheless, attention-related increases in sensory-evoked cortical activity were only observed at the parafoveal target location, as measured by the amplitude of the lateral occipital P1 ERP component. A second experiment replicated this data pattern using targets with lower stimulus contrast, indicating that the absence of a P1 effect for foveal targets could not be attributed to a saturated P1 response under higher-contrast stimulus conditions. When considered in light of retinogeniculate projections to cortex showing systematic changes in their physiological organization beginning within a degree of visual angle of the fovea, our findings support the proposal that the strategic functions of visual attention may vary with the retinotopic location involved.
Journal Articles
Michael B. Miller, John Darrell Van Horn, George L. Wolford, Todd C. Handy, Monica Valsangkar-Smyth ...
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2002) 14 (8): 1200–1214.
Published: 01 November 2002
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The localization of brain functions using neuroimaging techniques is commonly dependent on statistical analyses of groups of subjects in order to identify sites of activation, particularly in studies of episodic memory. Exclusive reliance on group analysis may be to the detriment of understanding the true underlying cognitive nature of brain activations. In the present study, we found that the patterns of brain activity associated with episodic retrieval are very distinct for individual subjects from the patterns of brain activity at the group level. These differences go beyond the relatively small variations due to cyctoarchitectonic differences or spatial normalization. We quantify this individual variability by cross-correlating volumes of brain images. We demonstrate that individual patterns of brain activity are reliable over time despite their extensive variability. We suggest that varied but reliable individual patterns of significant brain activity may be indicative of different cognitive strategies used to produce a recognition response. We believe that individual analysis in conjunction with group analysis may be critical to fully understanding the relationship between retrieval processes and underlying brain regions.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2000) 12 (6): 1066–1069.
Published: 01 November 2000
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Growing evidence suggests that a more complete understanding of cortical function requires developing cognitive models that are predictive of multivariate neural behavior (e.g., Raichle, 2000; Shulman et al., 1997). Towards this end, one approach is to interpret population-specific activity in cortex from the perspective of capacity theories of selective attention (e.g., Handy, Hopfinger, & Mangun, in press). In brief, the model is founded on the ideas that (1) processing capacity is limited and (2) different processes may draw on different capacities (e.g., Boles & Law 1998; Polson & Friedman, 1988; see Kramer & Spinks, 1991). Testable hypotheses are then based on whether manipulations of process-specific load will lead to negative or positive covariation between different function-related clusters of activation in cortex—the predicted pattern depends on whether or not the clusters in question are assumed to share a common processing capacity. Expanding on these ideas, the current article addresses several recent issues that have arisen in the effort to apply capacity theory to the study of cortical function.