Browsing by Subject "Reaction Time"

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    A neurophysiological study into the foundations of tonal harmony.
    (Neuroreport, 2009-02-18) Bergelson, Elika; Idsardi, William J
    Our findings provide magnetoencephalographic evidence that the mismatch-negativity response to two-note chords (dyads) is modulated by a combination of abstract cognitive differences and lower-level differences in the auditory signal. Participants were presented with series of simple-ratio sinusoidal dyads (perfect fourths and perfect fifths) in which the difference between the standard and deviant dyad exhibited an interval change, a shift in pitch space, or both. In addition, the standard-deviant pair of dyads either shared one note or both notes were changed. Only the condition that featured both abstract changes (interval change and pitch-space shift) and two novel notes showed a significantly larger magnetoencephalographic mismatch-negativity response than the other conditions in the right hemisphere. Implications for music and language processing are discussed.
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    A wireless multi-channel recording system for freely behaving mice and rats.
    (PLoS One, 2011) Fan, David; Rich, Dylan; Holtzman, Tahl; Ruther, Patrick; Dalley, Jeffrey W; Lopez, Alberto; Rossi, Mark A; Barter, Joseph W; Salas-Meza, Daniel; Herwik, Stanislav; Holzhammer, Tobias; Morizio, James; Yin, Henry H
    To understand the neural basis of behavior, it is necessary to record brain activity in freely moving animals. Advances in implantable multi-electrode array technology have enabled researchers to record the activity of neuronal ensembles from multiple brain regions. The full potential of this approach is currently limited by reliance on cable tethers, with bundles of wires connecting the implanted electrodes to the data acquisition system while impeding the natural behavior of the animal. To overcome these limitations, here we introduce a multi-channel wireless headstage system designed for small animals such as rats and mice. A variety of single unit and local field potential signals were recorded from the dorsal striatum and substantia nigra in mice and the ventral striatum and prefrontal cortex simultaneously in rats. This wireless system could be interfaced with commercially available data acquisition systems, and the signals obtained were comparable in quality to those acquired using cable tethers. On account of its small size, light weight, and rechargeable battery, this wireless headstage system is suitable for studying the neural basis of natural behavior, eliminating the need for wires, commutators, and other limitations associated with traditional tethered recording systems.
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    Adult age differences in functional connectivity during executive control.
    (Neuroimage, 2010-08-15) Madden, David J; Costello, Matthew C; Dennis, Nancy A; Davis, Simon W; Shepler, Anne M; Spaniol, Julia; Bucur, Barbara; Cabeza, Roberto
    Task switching requires executive control processes that undergo age-related decline. Previous neuroimaging studies have identified age-related differences in brain activation associated with global switching effects (dual-task blocks versus single-task blocks), but age-related differences in activation during local switching effects (switch trials versus repeat trials, within blocks) have not been investigated. This experiment used functional magnetic resonance imaging (fMRI), and diffusion tensor imaging (DTI), to examine adult age differences in task switching across adjacent trials (i.e., local task switching). During fMRI scanning, participants performed a cued, word categorization task. From interspersed cue-only trials, switch-related processing associated with the cue was estimated separately from the target. Activation associated with task switching, within a distributed frontoparietal network, differed for cue- and target-related processing. The magnitude of event-related activation for task switching was similar for younger adults (n=20; 18-27years) and older adults (n=20; 60-85years), although activation sustained throughout the on-tasks periods exhibited some age-related decline. Critically, the functional connectivity of switch-related regions, during cue processing, was higher for younger adults than for older adults, whereas functional connectivity during target processing was comparable across the age groups. Further, individual differences in cue-related functional connectivity shared a substantial portion of the age-related variability in the efficiency of target categorization response (drift rate). This age-related difference in functional connectivity, however, was independent of white matter integrity within task-relevant regions. These findings highlight the functional connectivity of frontoparietal activation as a potential source of age-related decline in executive control.
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    Age mediation of frontoparietal activation during visual feature search.
    (Neuroimage, 2014-11-15) Madden, David J; Parks, Emily L; Davis, Simon W; Diaz, Michele T; Potter, Guy G; Chou, Ying-hui; Chen, Nan-kuei; Cabeza, Roberto
    Activation of frontal and parietal brain regions is associated with attentional control during visual search. We used fMRI to characterize age-related differences in frontoparietal activation in a highly efficient feature search task, detection of a shape singleton. On half of the trials, a salient distractor (a color singleton) was present in the display. The hypothesis was that frontoparietal activation mediated the relation between age and attentional capture by the salient distractor. Participants were healthy, community-dwelling individuals, 21 younger adults (19-29 years of age) and 21 older adults (60-87 years of age). Top-down attention, in the form of target predictability, was associated with an improvement in search performance that was comparable for younger and older adults. The increase in search reaction time (RT) associated with the salient distractor (attentional capture), standardized to correct for generalized age-related slowing, was greater for older adults than for younger adults. On trials with a color singleton distractor, search RT increased as a function of increasing activation in frontal regions, for both age groups combined, suggesting increased task difficulty. Mediational analyses disconfirmed the hypothesized model, in which frontal activation mediated the age-related increase in attentional capture, but supported an alternative model in which age was a mediator of the relation between frontal activation and capture.
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    Age-related differences in frontoparietal activation for target and distractor singletons during visual search.
    (Attention, perception & psychophysics, 2023-04) Merenstein, Jenna L; Mullin, Hollie A; Madden, David J
    Age-related decline in visual search performance has been associated with different patterns of activation in frontoparietal regions using functional magnetic resonance imaging (fMRI), but whether these age-related effects represent specific influences of target and distractor processing is unclear. Therefore, we acquired event-related fMRI data from 68 healthy, community-dwelling adults ages 18-78 years, during both conjunction (T/F target among rotated Ts and Fs) and feature (T/F target among Os) search. Some displays contained a color singleton that could correspond to either the target or a distractor. A diffusion decision analysis indicated age-related increases in sensorimotor response time across all task conditions, but an age-related decrease in the rate of evidence accumulation (drift rate) was specific to conjunction search. Moreover, the color singleton facilitated search performance when occurring as a target and disrupted performance when occurring as a distractor, but only during conjunction search, and these effects were independent of age. The fMRI data indicated that decreased search efficiency for conjunction relative to feature search was evident as widespread frontoparietal activation. Activation within the left insula mediated the age-related decrease in drift rate for conjunction search, whereas this relation in the FEF and parietal cortex was significant only for individuals younger than 30 or 44 years, respectively. Finally, distractor singletons were associated with significant parietal activation, whereas target singletons were associated with significant frontoparietal deactivation, and this latter effect increased with adult age. Age-related differences in frontoparietal activation therefore reflect both the overall efficiency of search and the enhancement from salient targets.
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    Age-related differences in resolving semantic and phonological competition during receptive language tasks.
    (Neuropsychologia, 2016-12) Zhuang, Jie; Johnson, Micah A; Madden, David J; Burke, Deborah M; Diaz, Michele T
    Receptive language (e.g., reading) is largely preserved in the aging brain, and semantic processes in particular may continue to develop throughout the lifespan. We investigated the neural underpinnings of phonological and semantic retrieval in older and younger adults during receptive language tasks (rhyme and semantic similarity judgments). In particular, we were interested in the role of competition on language retrieval and varied the similarities between a cue, target, and distractor that were hypothesized to affect the mental process of competition. Behaviorally, all participants responded faster and more accurately during the rhyme task compared to the semantic task. Moreover, older adults demonstrated higher response accuracy than younger adults during the semantic task. Although there were no overall age-related differences in the neuroimaging results, an Age×Task interaction was found in left inferior frontal gyrus (IFG), with older adults producing greater activation than younger adults during the semantic condition. These results suggest that at lower levels of task difficulty, older and younger adults engaged similar neural networks that benefited behavioral performance. As task difficulty increased during the semantic task, older adults relied more heavily on largely left hemisphere language regions, as well as regions involved in perception and internal monitoring. Our results are consistent with the stability of language comprehension across the adult lifespan and illustrate how the preservation of semantic representations with aging may influence performance under conditions of increased task difficulty.
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    Age-related differences in the neural bases of phonological and semantic processes.
    (Journal of cognitive neuroscience, 2014-12) Diaz, Michele T; Johnson, Micah A; Burke, Deborah M; Madden, David J
    Changes in language functions during normal aging are greater for phonological compared with semantic processes. To investigate the behavioral and neural basis for these age-related differences, we used fMRI to examine younger and older adults who made semantic and phonological decisions about pictures. The behavioral performance of older adults was less accurate and less efficient than younger adults' in the phonological task but did not differ in the semantic task. In the fMRI analyses, the semantic task activated left-hemisphere language regions, and the phonological task activated bilateral cingulate and ventral precuneus. Age-related effects were widespread throughout the brain and most often expressed as greater activation for older adults. Activation was greater for younger compared with older adults in ventral brain regions involved in visual and object processing. Although there was not a significant Age × Condition interaction in the whole-brain fMRI results, correlations examining the relationship between behavior and fMRI activation were stronger for younger compared with older adults. Our results suggest that the relationship between behavior and neural activation declines with age, and this may underlie some of the observed declines in performance.
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    Age-related effects on the neural correlates of autobiographical memory retrieval.
    (Neurobiol Aging, 2012-07) St Jacques, Peggy L; Rubin, David C; Cabeza, Roberto
    Older adults recall less episodically rich autobiographical memories (AM), however, the neural basis of this effect is not clear. Using functional MRI, we examined the effects of age during search and elaboration phases of AM retrieval. Our results suggest that the age-related attenuation in the episodic richness of AMs is associated with difficulty in the strategic retrieval processes underlying recovery of information during elaboration. First, age effects on AM activity were more pronounced during elaboration than search, with older adults showing less sustained recruitment of the hippocampus and ventrolateral prefrontal cortex (VLPFC) for less episodically rich AMs. Second, there was an age-related reduction in the modulation of top-down coupling of the VLPFC on the hippocampus for episodically rich AMs. In sum, the present study shows that changes in the sustained response and coupling of the hippocampus and prefrontal cortex (PFC) underlie age-related reductions in episodic richness of the personal past.
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    Age-related preservation of top-down control over distraction in visual search.
    (Experimental aging research, 2010-07) Costello, Matthew C; Madden, David J; Shepler, Anne M; Mitroff, Stephen R; Leber, Andrew B
    Visual search studies have demonstrated that older adults can have preserved or even increased top-down control over distraction. However, the results are mixed as to the extent of this age-related preservation. The present experiment assesses group differences in younger and older adults during visual search, with a task featuring two conditions offering varying degrees of top-down control over distraction. After controlling for generalized slowing, the analyses revealed that the age groups were equally capable of utilizing top-down control to minimize distraction. Furthermore, for both age groups, the distraction effect was manifested in a sustained manner across the reaction time distribution.
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    Age-related slowing in the retrieval of information from long-term memory.
    (Journal of gerontology, 1985-03) Madden, DJ
    The present experiment investigated adult age differences in the retrieval of information from long-term memory. Each trial required a decision regarding the synonymy of two visually presented words. On the yes-response trials, the two words were either identical, differed only in case, or were synonyms that differed in case. Age differences in absolute decision time were greater for the synonyms than for the other word pairs, but the proportional slowing of decision time exhibited by the older adults was constant across word-pair type. A generalized age-related slowing in the speed of information processing can currently account for age differences in the retrieval of letter-identity and semantic information from long-term memory.
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    Co-activation of the amygdala, hippocampus and inferior frontal gyrus during autobiographical memory retrieval.
    (Neuropsychologia, 2005) Greenberg, Daniel L; Rice, Heather J; Cooper, Julie J; Cabeza, Roberto; Rubin, David C; Labar, Kevin S
    Functional MRI was used to investigate the role of medial temporal lobe and inferior frontal lobe regions in autobiographical recall. Prior to scanning, participants generated cue words for 50 autobiographical memories and rated their phenomenological properties using our autobiographical memory questionnaire (AMQ). During scanning, the cue words were presented and participants pressed a button when they retrieved the associated memory. The autobiographical retrieval task was interleaved in an event-related design with a semantic retrieval task (category generation). Region-of-interest analyses showed greater activation of the amygdala, hippocampus, and right inferior frontal gyrus during autobiographical retrieval relative to semantic retrieval. In addition, the left inferior frontal gyrus showed a more prolonged duration of activation in the semantic retrieval condition. A targeted correlational analysis revealed pronounced functional connectivity among the amygdala, hippocampus, and right inferior frontal gyrus during autobiographical retrieval but not during semantic retrieval. These results support theories of autobiographical memory that hypothesize co-activation of frontotemporal areas during recollection of episodes from the personal past.
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    Composition and topographic organization of signals sent from the frontal eye field to the superior colliculus.
    (J Neurophysiol, 2000-04) Sommer, MA; Wurtz, RH
    The frontal eye field (FEF) and superior colliculus (SC) contribute to saccadic eye movement generation, and much of the FEF's oculomotor influence may be mediated through the SC. The present study examined the composition and topographic organization of signals flowing from FEF to SC by recording from FEF neurons that were antidromically activated from rostral or caudal SC. The first and most general result was that, in a sample of 88 corticotectal neurons, the types of signals relayed from FEF to SC were highly diverse, reflecting the general population of signals within FEF rather than any specific subset of signals. Second, many neurons projecting from FEF to SC carried signals thought to reflect cognitive operations, namely tonic discharges during the delay period of a delayed-saccade task (delay signals), elevated discharges during the gap period of a gap task (gap increase signals), or both. Third, FEF neurons discharging during fixation were found to project to the SC, although they did not project preferentially to rostral SC, where similar fixation neurons are found. Neurons that did project preferentially to the rostral SC were those with foveal visual responses and those pausing during the gap period of the gap task. Many of the latter neurons also had foveal visual responses, presaccadic pauses in activity, and postsaccadic increases in activity. These two types of rostral-projecting neurons therefore may contribute to the activity of rostral SC fixation neurons. Fourth, conduction velocity was used as an indicator of cell size to correct for sampling bias. The outcome of this correction procedure suggested that among the most prevalent neurons in the FEF corticotectal population are those carrying putative cognitive-related signals, i.e., delay and gap increase signals, and among the least prevalent are those carrying presaccadic burst discharges but lacking peripheral visual responses. Fifth, corticotectal neurons carrying various signals were biased topographically across the FEF. Neurons with peripheral visual responses but lacking presaccadic burst discharges were biased laterally, neurons with presaccadic burst discharges but lacking peripheral visual responses were biased medially, and neurons carrying delay or gap increase signals were biased dorsally. Finally, corticotectal neurons were distributed within the FEF as a function of their visual or movement field eccentricity and projected to the SC such that eccentricity maps in both structures were closely aligned. We conclude that the FEF most likely influences the activity of SC neurons continuously from the start of fixation, through visual analysis and cognitive manipulations, until a saccade is generated and fixation begins anew. Furthermore, the projection from FEF to SC is highly topographically organized in terms of function at both its source and its termination.
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    Cortical Brain Activity Reflecting Attentional Biasing Toward Reward-Predicting Cues Covaries with Economic Decision-Making Performance.
    (Cereb Cortex, 2016-01) San Martín, René; Appelbaum, Lawrence G; Huettel, Scott A; Woldorff, Marty G
    Adaptive choice behavior depends critically on identifying and learning from outcome-predicting cues. We hypothesized that attention may be preferentially directed toward certain outcome-predicting cues. We studied this possibility by analyzing event-related potential (ERP) responses in humans during a probabilistic decision-making task. Participants viewed pairs of outcome-predicting visual cues and then chose to wager either a small (i.e., loss-minimizing) or large (i.e., gain-maximizing) amount of money. The cues were bilaterally presented, which allowed us to extract the relative neural responses to each cue by using a contralateral-versus-ipsilateral ERP contrast. We found an early lateralized ERP response, whose features matched the attention-shift-related N2pc component and whose amplitude scaled with the learned reward-predicting value of the cues as predicted by an attention-for-reward model. Consistently, we found a double dissociation involving the N2pc. Across participants, gain-maximization positively correlated with the N2pc amplitude to the most reliable gain-predicting cue, suggesting an attentional bias toward such cues. Conversely, loss-minimization was negatively correlated with the N2pc amplitude to the most reliable loss-predicting cue, suggesting an attentional avoidance toward such stimuli. These results indicate that learned stimulus-reward associations can influence rapid attention allocation, and that differences in this process are associated with individual differences in economic decision-making performance.
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    Cross-hemispheric collaboration and segregation associated with task difficulty as revealed by structural and functional connectivity.
    (J Neurosci, 2015-05-27) Davis, Simon W; Cabeza, Roberto
    Although it is known that brain regions in one hemisphere may interact very closely with their corresponding contralateral regions (collaboration) or operate relatively independent of them (segregation), the specific brain regions (where) and conditions (how) associated with collaboration or segregation are largely unknown. We investigated these issues using a split field-matching task in which participants matched the meaning of words or the visual features of faces presented to the same (unilateral) or to different (bilateral) visual fields. Matching difficulty was manipulated by varying the semantic similarity of words or the visual similarity of faces. We assessed the white matter using the fractional anisotropy (FA) measure provided by diffusion tensor imaging (DTI) and cross-hemispheric communication in terms of fMRI-based connectivity between homotopic pairs of cortical regions. For both perceptual and semantic matching, bilateral trials became faster than unilateral trials as difficulty increased (bilateral processing advantage, BPA). The study yielded three novel findings. First, whereas FA in anterior corpus callosum (genu) correlated with word-matching BPA, FA in posterior corpus callosum (splenium-occipital) correlated with face-matching BPA. Second, as matching difficulty intensified, cross-hemispheric functional connectivity (CFC) increased in domain-general frontopolar cortex (for both word and face matching) but decreased in domain-specific ventral temporal lobe regions (temporal pole for word matching and fusiform gyrus for face matching). Last, a mediation analysis linking DTI and fMRI data showed that CFC mediated the effect of callosal FA on BPA. These findings clarify the mechanisms by which the hemispheres interact to perform complex cognitive tasks.
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    Cross-modal stimulus conflict: the behavioral effects of stimulus input timing in a visual-auditory Stroop task.
    (PLoS One, 2013) Donohue, Sarah E; Appelbaum, Lawrence G; Park, Christina J; Roberts, Kenneth C; Woldorff, Marty G
    Cross-modal processing depends strongly on the compatibility between different sensory inputs, the relative timing of their arrival to brain processing components, and on how attention is allocated. In this behavioral study, we employed a cross-modal audio-visual Stroop task in which we manipulated the within-trial stimulus-onset-asynchronies (SOAs) of the stimulus-component inputs, the grouping of the SOAs (blocked vs. random), the attended modality (auditory or visual), and the congruency of the Stroop color-word stimuli (congruent, incongruent, neutral) to assess how these factors interact within a multisensory context. One main result was that visual distractors produced larger incongruency effects on auditory targets than vice versa. Moreover, as revealed by both overall shorter response times (RTs) and relative shifts in the psychometric incongruency-effect functions, visual-information processing was faster and produced stronger and longer-lasting incongruency effects than did auditory. When attending to either modality, stimulus incongruency from the other modality interacted with SOA, yielding larger effects when the irrelevant distractor occurred prior to the attended target, but no interaction with SOA grouping. Finally, relative to neutral-stimuli, and across the wide range of the SOAs employed, congruency led to substantially more behavioral facilitation than did incongruency to interference, in contrast to findings that within-modality stimulus-compatibility effects tend to be more evenly split between facilitation and interference. In sum, the present findings reveal several key characteristics of how we process the stimulus compatibility of cross-modal sensory inputs, reflecting stimulus processing patterns that are critical for successfully navigating our complex multisensory world.
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    Data-driven and memory-driven selective attention in visual search.
    (Journal of gerontology, 1984-01) Madden, DJ
    The present experiment investigated Rabbitt's (1979) hypothesis that age differences in selective attention occur when memory-driven processing must be employed. Young and older adults performed a visual search task, which, on some trials, provided advance information (a cue) regarding the particular target letter most likely to appear in the display. The nature of the selectivity required by the cue was either data-driven (Condition 1) or memory-driven (Condition 2). Analyses of the benefit in search performance associated with the cued trials and of the cost in performance resulting from the presentation of misleading advance information yielded limited support for Rabbitt's hypothesis. The older adults, but not the young, did exhibit a smaller cuing benefit in Condition 2 than in Condition 1. Both age groups, however, demonstrated substantial benefits and costs within each condition. Age differences in selective attention are thus not determined completely by the requirement to use memory-driven processing.
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    Delay activity of saccade-related neurons in the caudal dentate nucleus of the macaque cerebellum.
    (J Neurophysiol, 2013-04) Ashmore, Robin C; Sommer, Marc A
    The caudal dentate nucleus (DN) in lateral cerebellum is connected with two visual/oculomotor areas of the cerebrum: the frontal eye field and lateral intraparietal cortex. Many neurons in frontal eye field and lateral intraparietal cortex produce "delay activity" between stimulus and response that correlates with processes such as motor planning. Our hypothesis was that caudal DN neurons would have prominent delay activity as well. From lesion studies, we predicted that this activity would be related to self-timing, i.e., the triggering of saccades based on the internal monitoring of time. We recorded from neurons in the caudal DN of monkeys (Macaca mulatta) that made delayed saccades with or without a self-timing requirement. Most (84%) of the caudal DN neurons had delay activity. These neurons conveyed at least three types of information. First, their activity was often correlated, trial by trial, with saccade initiation. Correlations were found more frequently in a task that required self-timing of saccades (53% of neurons) than in a task that did not (27% of neurons). Second, the delay activity was often tuned for saccade direction (in 65% of neurons). This tuning emerged continuously during a trial. Third, the time course of delay activity associated with self-timed saccades differed significantly from that associated with visually guided saccades (in 71% of neurons). A minority of neurons had sensory-related activity. None had presaccadic bursts, in contrast to DN neurons recorded more rostrally. We conclude that caudal DN neurons convey saccade-related delay activity that may contribute to the motor preparation of when and where to move.
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    Development of hemispheric specialization for lexical pitch-accent in Japanese infants.
    (J Cogn Neurosci, 2010-11) Sato, Yutaka; Sogabe, Yuko; Mazuka, Reiko
    Infants' speech perception abilities change through the first year of life, from broad sensitivity to a wide range of speech contrasts to becoming more finely attuned to their native language. What remains unclear, however, is how this perceptual change relates to brain responses to native language contrasts in terms of the functional specialization of the left and right hemispheres. Here, to elucidate the developmental changes in functional lateralization accompanying this perceptual change, we conducted two experiments on Japanese infants using Japanese lexical pitch-accent, which changes word meanings with the pitch pattern within words. In the first behavioral experiment, using visual habituation, we confirmed that infants at both 4 and 10 months have sensitivities to the lexical pitch-accent pattern change embedded in disyllabic words. In the second experiment, near-infrared spectroscopy was used to measure cortical hemodynamic responses in the left and right hemispheres to the same lexical pitch-accent pattern changes and their pure tone counterparts. We found that brain responses to the pitch change within words differed between 4- and 10-month-old infants in terms of functional lateralization: Left hemisphere dominance for the perception of the pitch change embedded in words was seen only in the 10-month-olds. These results suggest that the perceptual change in Japanese lexical pitch-accent may be related to a shift in functional lateralization from bilateral to left hemisphere dominance.
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    Effects of adult age and blood pressure on executive function and speed of processing.
    (Experimental aging research, 2010-04) Bucur, Barbara; Madden, David J
    Previous research has established that the effects of chronically increased blood pressure (BP) on cognition interact with adult age, but the relevant cognitive processes are not well defined. In this cross-sectional study, using a sample matched for age, years of education, and sex, 134 individuals with either normal BP (n = 71) or chronically high BP (n = 63) were categorized into younger (19-39 years), middle-aged (41-58 years), and older (60-79 years) groups. Using a between-subjects analysis of variance (ANOVA), covarying for race and years of education, composite measures of executive function and perceptual speed both exhibited age-related decline. The executive function measure, however, was associated with a differential decline in high BP older adults. This result held even when the executive function scores were covaried for speed, demonstrating an independent, age-related effect of higher BP on executive function.
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    Express averaging saccades in monkeys.
    (Vision Res, 1999) Chou, IH; Sommer, MA; Schiller, PH
    When monkeys are presented simultaneously with multiple stimuli, they can make one of two types of response. Either they make averaging saccades, that land at intermediate locations between the targets, or target-directed saccades, that land close to one of the targets. The two types of saccades occur at different latencies and are thought to reflect different processes; fast reflexive averaging and slower target selection. We investigated the latency of averaging saccades in five monkeys, with particular emphasis on 'express' latency saccades, which are thought to be inhibited by target selection. Express averaging saccades were made prolifically by the two monkeys that made both express and regular latency saccades, but only when no specific instruction was given regarding the saccade target. When these monkeys had to choose one of the targets, on the basis of its color, they still made averaging saccades. However, the endpoints formed two distributions close to the targets as opposed to one single distribution centered between the targets, as was the case when targets were identical; also, express saccades were almost entirely absent. We conclude that express averaging saccades are a form of spatial and temporal optimization of gaze shifting.