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1.  Faces are special but not too special: Spared face recognition in amnesia is based on familiarity 
Neuropsychologia  2010;48(13):3941-3948.
Most current theories of human memory are material-general in the sense that they assume that the medial temporal lobe (MTL) is important for retrieving the details of prior events, regardless of the specific type of materials. Recent studies of amnesia have challenged the material-general assumption by suggesting that the MTL may be necessary for remembering words, but is not involved in remembering faces. We examined recognition memory for faces and words in a group of amnesic patients, which included hypoxic patients and patients with extensive left or right MTL lesions. Recognition confidence judgments were used to plot receiver operating characteristics (ROCs) in order to more fully quantify recognition performance and to estimate the contributions of recollection and familiarity. Consistent with the extant literature, an analysis of overall recognition accuracy showed that the patients were impaired at word memory but had spared face memory. However, the ROC analysis indicated that the patients were generally impaired at high confidence recognition responses for faces and words, and they exhibited significant recollection impairments for both types of materials. Familiarity for faces was preserved in all patients, but extensive left MTL damage impaired familiarity for words. These results suggest that face recognition may appear to be spared because performance tends to rely heavily on familiarity, a process that is relatively well preserved in amnesia. The findings challenge material-general theories of memory, and suggest that both material and process are important determinants of memory performance in amnesia, and different types of materials may depend more or less on recollection and familiarity.
doi:10.1016/j.neuropsychologia.2010.09.005
PMCID: PMC4084520  PMID: 20833190
episodic memory; recollection; familiarity; amnesia; face recognition; word recognition
2.  Stimulation of contacts in ventral but not dorsal subthalamic nucleus normalizes response switching in Parkinson's disease 
Neuropsychologia  2013;51(7):1302-1309.
Switching between responses is a key executive function known to rely on the frontal cortex and the basal ganglia. Here we aimed to establish with greater anatomical specificity whether such switching could be mediated via different possible frontal–basal-ganglia circuits. Accordingly, we stimulated dorsal vs. ventral contacts of electrodes in the subthalamic nucleus (STN) in Parkinson's patients during switching performance, and also studied matched controls. The patients underwent three sessions: once with bilateral dorsal contact stimulation, once with bilateral ventral contact stimulation, and once Off stimulation. Patients Off stimulation showed abnormal patterns of switching, and stimulation of the ventral contacts but not the dorsal contacts normalized the pattern of behavior relative to controls. This provides some of the first evidence in humans that stimulation of dorsal vs. ventral STN DBS contacts has differential effects on executive function. As response switching is an executive function known to rely on prefrontal cortex, these results suggest that ventral contact stimulation affected an executive/associative cortico-basal ganglia circuit.
doi:10.1016/j.neuropsychologia.2013.03.008
PMCID: PMC4079723  PMID: 23562963
Cognitive control; Switching; Subthalamic nucleus; Deep brain stimulation; Parkinson's disease
3.  Naming and gesturing spatial relations: Evidence from focal brain-injured individuals 
Neuropsychologia  2013;51(8):1518-1527.
Spatial language helps us encode relations between objects and organize our thinking. Little is known about the neural instantiations of spatial language. Using voxel-lesion symptom mapping (VLSM), we tested the hypothesis that focal brain injured patients who had damage to left frontal-parietal peri-Sylvian regions would have difficulty in naming spatial relations between objects. We also investigated the relationship between impaired verbalization of spatial relations and spontaneous gesture production. Patients with left or right hemisphere damage and elderly control participants were asked to name static (e.g., an apple on a book) and dynamic (e.g., a pen moves over a box) locative relations depicted in brief video clips. The correct use of prepositions in each task and gestures that represent the spatial relations were coded. Damage to the left posterior middle frontal gyrus, the left inferior frontal gyrus, and the left anterior superior temporal gyrus were related to impairment in naming spatial relations. Production of spatial gestures negatively correlated with naming accuracy, suggesting that gestures might help or compensate for difficulty with lexical access. Additional analyses suggested that left hemisphere patients who had damage to the left posterior middle frontal gyrus and the left inferior frontal gyrus gestured less than expected, if gestures are used to compensate for impairments in retrieving prepositions.
doi:10.1016/j.neuropsychologia.2013.05.006
PMCID: PMC3699899  PMID: 23685196
lesion studies; spatial relations; prepositions; co-speech gestures
4.  Modulation of effective connectivity during vocalization with perturbed auditory feedback 
Neuropsychologia  2013;51(8):1471-1480.
The integration of auditory feedback with vocal motor output is important for the control of voice fundamental frequency (F0). We used a pitch-shift paradigm where subjects respond to an alteration, or shift, of voice pitch auditory feedback with a reflexive change in F0. We presented varying magnitudes of pitch shifted auditory feedback to subjects during vocalization and passive listening and measured event related potentials (ERP’s) to the feedback shifts. Shifts were delivered at +100 and +400 cents (200 ms duration). The ERP data were modeled with Dynamic Causal Modeling (DCM) techniques where the effective connectivity between the superior temporal gyrus (STG), inferior frontal gyrus and premotor areas were tested. We compared three main factors; the effect of intrinsic STG connectivity, STG modulation across hemispheres and the specific effect of hemisphere. A Bayesian model selection procedure was used to make inference about model families. Results suggest that both intrinsic STG and left to right STG connections are important in the identification of self-voice error and sensory motor integration. We identified differences in left to right STG connections between 100 cent and 400 cent shift conditions suggesting that self and non-self voice error are processed differently in the left and right hemisphere. These results also highlight the potential of DCM modeling of ERP responses to characterize specific network properties of forward models of voice control.
doi:10.1016/j.neuropsychologia.2013.05.002
PMCID: PMC3704150  PMID: 23665378
Vocalization; Auditory feedback; ERP; DCM; Audio-vocal integration; Pitch shift
5.  Prefrontal vulnerabilities and whole brain connectivity in aging and depression 
Neuropsychologia  2013;51(8):1463-1470.
Studies exploring the underpinnings of age-related neurodegeneration suggest fronto-limbic alterations that are increasingly vulnerable in the presence of disease including late life depression. Less work has assessed the impact of this specific vulnerability on widespread brain circuitry. Seventy-nine older adults (healthy controls=45; late life depression=34) completed translational tasks shown in non-human primates to rely on fronto-limbic networks involving dorsolateral (Self-Ordered Pointing Task) or orbitofrontal (Object Alternation Task) cortices. A sub-sample of participants also completed diffusion tensor imaging for white matter tract quantification (uncinate and cingulum bundle; n=58) and whole brain tract-based spatial statistics (n=62). Despite task associations to specific white matter tracts across both groups, only healthy controls demonstrated significant correlations between widespread tract integrity and cognition. Thus, increasing Object Alternation Task errors were associated with decreasing fractional anisotropy in the uncinate in late life depression; however, only in healthy controls was the uncinate incorporated into a larger network of white matter vulnerability associating fractional anisotropy with Object Alternation Task errors using whole brain tract-based spatial statistics. It appears that the whole brain impact of specific fronto-limbic vulnerabilities in aging may be eclipsed in the presence of disease-specific neuropathology like that seen in late life depression.
doi:10.1016/j.neuropsychologia.2013.05.004
PMCID: PMC3726267  PMID: 23680399
ageing; prefrontal vulnerability; white matter tracts; myelin integrity; late life depression
6.  Where is the action? Action sentence processing in Parkinson's disease 
Neuropsychologia  2013;51(8):1510-1517.
According to an influential view of conceptual representation, action concepts are understood through motoric simulations, involving motor networks of the brain. A stronger version of this embodied account suggests that even figurative uses of action words (e.g., grasping the concept) are understood through motoric simulations. We investigated these claims by assessing whether Parkinson's disease (PD), a disorder affecting the motor system, is associated with selective deficits in comprehending action-related sentences. Twenty PD patients and 21 age-matched controls performed a sentence comprehension task, where sentences belonged to one of four conditions: literal action, non-idiomatic metaphoric action, idiomatic action, and abstract. The same verbs (referring to hand/arm actions) were used in the three action-related conditions. Patients, but not controls, were slower to respond to literal and idiomatic action than to abstract sentences. These results indicate that sensory-motor systems play a functional role in semantic processing, including processing of figurative action language.
doi:10.1016/j.neuropsychologia.2013.04.008
PMCID: PMC3731159  PMID: 23624313
Conceptual processing; Embodiment; Figurative language; Language comprehension; Metaphor; Idiom; Sentence processing; Parkinson's disease
7.  Brain Function Overlaps When People Observe Emblems, Speech, and Grasping 
Neuropsychologia  2013;51(8):10.1016/j.neuropsychologia.2013.03.022.
A hand grasping a cup or gesturing ‘thumbs-up’, while both manual actions, have different purposes and effects. Grasping directly affects the cup, whereas gesturing ‘thumbs-up’ has an effect through an implied verbal (symbolic) meaning. Because grasping and emblematic gestures (‘emblems’) are both goal-oriented hand actions, we pursued the hypothesis that observing each should evoke similar activity in neural regions implicated in processing goal-oriented hand actions. However, because emblems express symbolic meaning, observing them should also evoke activity in regions implicated in interpreting meaning, which is most commonly expressed in language. Using fMRI to test this hypothesis, we had participants watch videos of an actor performing emblems, speaking utterances matched in meaning to the emblems, and grasping objects. Our results show that lateral temporal and inferior frontal regions respond to symbolic meaning, even when it is expressed by a single hand action. In particular, we found that left inferior frontal and right lateral temporal regions are strongly engaged when people observe either emblems or speech. In contrast, we also replicate and extend previous work that implicates parietal and premotor responses in observing goal-oriented hand actions. For hand actions, we found that bilateral parietal and premotor regions are strongly engaged when people observe either emblems or grasping. These findings thus characterize converging brain responses to shared features (e.g., symbolic or manual), despite their encoding and presentation in different stimulus modalities.
doi:10.1016/j.neuropsychologia.2013.03.022
PMCID: PMC3813464  PMID: 23583968
gestures; language; semantics; perception; functional magnetic resonance imaging
8.  The association between higher order abilities, processing speed, and age are variably mediated by white matter integrity during typical aging 
Neuropsychologia  2013;51(8):10.1016/j.neuropsychologia.2013.03.005.
Although aging is associated with changes in brain structure and cognition it remains unclear which specific structural changes mediate individual cognitive changes. Several studies have reported that white matter (WM) integrity, as assessed by diffusion tensor imaging (DTI), mediates, in part, age-related differences in processing speed (PS). There is less evidence for WM integrity mediating age-related differences in higher order abilities (e.g., memory and executive functions). In 165 typically aging adults (age range 54–89) we show that WM integrity in select cerebral regions is associated with higher cognitive abilities and accounts variance not accounted for by PS or age. Specifically, voxel-wise analyses using tract-based spatial statistics (TBSS) revealed that WM integrity was associated with reasoning, cognitive flexibility and PS, but not memory or word fluency, after accounting for age and gender. While cerebral fractional anisotropy (FA) was only associated with PS; mean (MD), axial (AD) and radial (RD) diffusivity were associated with reasoning and flexibility. Reasoning was selectively associated with left prefrontal AD, while cognitive flexibility was associated with MD, AD and RD throughout the cerebrum. Average WM metrics within select WM regions of interest accounted for 18% and 29% of the variance in reasoning and flexibility, respectively, similar to the amount of variance accounted for by age. WM metrics mediated ~50% of the age-related variance in reasoning and flexibility and different proportions, 11% for reasoning and 44% for flexibility, of the variance accounted for by PS. In sum, i) WM integrity is significantly, but variably, related to specific higher cognitive abilities and can account for a similar proportion of variance as age, and ii) while FA is selectively associated with PS; while MD, AD and RD are associated with reasoning, flexibility and PS. This illustrates both the anatomical and cognitive selectivity of structure-cognition relationships in the aging brain.
doi:10.1016/j.neuropsychologia.2013.03.005
PMCID: PMC3875161  PMID: 23507612
DTI; aging; cognition; MRI; processing speed; executive function; TBSS
9.  An electrophysiological index of changes in risk decision-making strategies 
Neuropsychologia  2013;51(8):1397-1407.
Human decision-making is significantly modulated by previously experienced outcomes. Using event-related potentials (ERPs), we examined whether ERP components evoked by outcome feedbacks could serve as biomarkers to signal the influence of current outcome evaluation on subsequent decision-making. In this study, eighteen adult volunteers participated in a simple monetary gambling task, in which they were asked to choose between two options that differed in risk. Their decisions were immediately followed by outcome presentation. Temporospatial principle component analysis (PCA) was applied to the outcome-onset locked ERPs in the -200 – 1000 ms time window. The PCA factors that approximated classical ERP components (P2, feedback-related negativity, P3a, & P3b) in terms of time course and scalp distribution were tested for their association with subsequent decision-making strategies. Our results revealed that a fronto-central PCA factor approximating the classical P3a was related to changes of decision-making strategies on subsequent trials. The decision to switch between high- and low-risk options resulted in a larger P3a relative to the decision to retain the same choice. According to the results, we suggest the amplitude of the fronto-central P3a is an electrophysiological index of the influence of current outcome on subsequent risk decision-making. Furthermore, the ERP source analysis indicated that the activations of the frontopolar cortex and sensorimotor cortex were involved in subsequent changes of strategies, which enriches our understanding of the neural mechanisms of adjusting decision-making strategies based on previous experience.
doi:10.1016/j.neuropsychologia.2013.04.014
PMCID: PMC3989206  PMID: 23643796
decision-making; outcome evaluation; event-related potential (ERP); principal components analysis (PCA); P3 component
10.  Speech comprehension aided by multiple modalities: behavioural and neural interactions 
Neuropsychologia  2012;50(5):762-776.
Speech comprehension is a complex human skill, the performance of which requires the perceiver to combine information from several sources – e.g. voice, face, gesture, linguistic context – to achieve an intelligible and interpretable percept. We describe a functional imaging investigation of how auditory, visual and linguistic information interact to facilitate comprehension. Our specific aims were to investigate the neural responses to these different information sources, alone and in interaction, and further to use behavioural speech comprehension scores to address sites of intelligibility-related activation in multifactorial speech comprehension. In fMRI, participants passively watched videos of spoken sentences, in which we varied Auditory Clarity (with noise-vocoding), Visual Clarity (with Gaussian blurring) and Linguistic Predictability. Main effects of enhanced signal with increased auditory and visual clarity were observed in overlapping regions of posterior STS. Two-way interactions of the factors (auditory × visual, auditory × predictability) in the neural data were observed outside temporal cortex, where positive signal change in response to clearer facial information and greater semantic predictability was greatest at intermediate levels of auditory clarity. Overall changes in stimulus intelligibility by condition (as determined using an independent behavioural experiment) were reflected in the neural data by increased activation predominantly in bilateral dorsolateral temporal cortex, as well as inferior frontal cortex and left fusiform gyrus. Specific investigation of intelligibility changes at intermediate auditory clarity revealed a set of regions, including posterior STS and fusiform gyrus, showing enhanced responses to both visual and linguistic information. Finally, an individual differences analysis showed that greater comprehension performance in the scanning participants (measured in a post-scan behavioural test) were associated with increased activation in left inferior frontal gyrus and left posterior STS. The current multimodal speech comprehension paradigm demonstrates recruitment of a wide comprehension network in the brain, in which posterior STS and fusiform gyrus form sites for convergence of auditory, visual and linguistic information, while left-dominant sites in temporal and frontal cortex support successful comprehension.
doi:10.1016/j.neuropsychologia.2012.01.010
PMCID: PMC4050300  PMID: 22266262
speech; fMRI; auditory cortex; individual differences; noise-vocoding
11.  Distinct critical cerebellar subregions for components of verbal working memory 
Neuropsychologia  2011;50(1):189-197.
A role for the cerebellum in cognition has been proposed based on studies suggesting a profile of cognitive deficits due to cerebellar stroke. Such studies are limited in the determination of the detailed organisation of cerebellar subregions that are critical for different aspects of cognition. In this study we examined the correlation between cognitive performance and cerebellar integrity in a specific degeneration of the cerebellar cortex: Spinocerebellar Ataxia type 6 (SCA6). The results demonstrate a critical relationship between verbal working memory and grey matter density in superior (bilateral lobules VI and crus I of lobule VII) and inferior (bilateral lobules VIIIa and VIIIb, and right lobule IX) parts of the cerebellum. We demonstrate that distinct cerebellar regions subserve different components of the prevalent psychological model for verbal working memory based on a phonological loop. The work confirms the involvement of the cerebellum in verbal working memory and defines specific subsystems for this within the cerebellum.
doi:10.1016/j.neuropsychologia.2011.11.017
PMCID: PMC4040406  PMID: 22133495
SCA-6; Cerebellum; Cognition; MRI; VBM; Neurodegeneration
12.  Dorsolateral Prefrontal Contributions to Human Intelligence 
Neuropsychologia  2012;51(7):1361-1369.
Although cognitive neuroscience has made remarkable progress in understanding the involvement of the prefrontal cortex in executive control functions for human intelligence, the necessity of the dorsolateral prefrontal cortex (dlPFC) for key competencies of general intelligence and executive function remains to be well established. Here we studied human brain lesion patients with dlPFC lesions to investigate whether this region is computationally necessary for performance on neuropsychological tests of general intelligence and executive function, administering the Wechsler Adult Intelligence Scale (WAIS) and subtests of the Delis Kaplan Executive Function System (D-KEFS) to three groups: dlPFC lesions (n = 19), non-dlPFC lesions (n = 152), and no brain lesions (n = 55). The key results indicate that: (1) patients with focal dlPFC damage exhibit lower scores, at the latent variable level, than controls in general intelligence (g) and executive function; (2) dlPFC patients demonstrate lower scores than controls in several executive measures; and (3) these latter differences are no longer significant when the pervasive influence of the general factor of intelligence (g) is statistically removed. The observed findings support a central role for the dlPFC in general intelligence and make specific recommendations for the interpretation and application of the WAIS and D-KEFS to the study of high-level cognition in health and disease.
doi:10.1016/j.neuropsychologia.2012.05.017
PMCID: PMC3478435  PMID: 22634247
prefrontal cortex; dorsolateral prefrontal cortex; general intelligence; executive function; lesion evidence
13.  Dissociations of action means and outcome processing in left hemisphere stroke 
Neuropsychologia  2013;51(7):1224-1233.
Previous evidence suggests that distinct fronto-parietal regions may be involved in representing action kinematics (means) and action results (outcome) during action observation. However, the evidence is contradictory with respect to the precise regions that are critical for each type of representation. Additionally unknown is the degree to which ability to detect action means and outcome during observation is related to action production performance. We used a behavioral task to evaluate the ability of healthy and left-hemisphere stroke participants to detect differences between pairs of videos that dissociated object-related action means (e.g., wiping with circular or straight movement) and/or outcome (e.g., applying or removing detergent). We expected that deficits in detecting action means would be associated with spatiomotor gesture production deficits, whereas deficits in detecting action outcome would predict impairments in complex naturalistic action. We also hypothesized a posterior to anterior gradient in the regions critical for each type of representation, disproportionately affecting means and outcome encoding, respectively. Results indicated that outcome – but not means – detection predicted naturalistic action performance in stroke participants. Regression and voxel lesion-symptom mapping analyses of lesion data revealed that means – but not outcome – coding relies on the integrity of the left inferior parietal lobe, whereas no selective critical brain region could be identified for outcome detection. Thus, means and outcome representations are dissociable at both the behavioral and neuroanatomical levels. Furthermore, the data are consistent with a degree of parallelism between action perception and production tasks. Finally, they reinforce the evidence for a critical role of the left inferior parietal lobule in the representation of action means, whereas action outcome may rely on a more distributed neural circuit.
doi:10.1016/j.neuropsychologia.2013.03.017
PMCID: PMC3690376  PMID: 23566892
Action representation; Means; Outcome; Stroke; Object-related action; Voxel-based lesion-symptom mapping
14.  Adaptive significance of right hemisphere activation in aphasic language comprehension 
Neuropsychologia  2013;51(7):10.1016/j.neuropsychologia.2013.03.007.
Aphasic patients often exhibit increased right hemisphere activity during language tasks. This may represent takeover of function by regions homologous to the left-hemisphere language networks, maladaptive interference, or adaptation of alternate compensatory strategies. To distinguish between these accounts, we tested language comprehension in 25 aphasic patients using an online sentence-picture matching paradigm while measuring brain activation with MEG. Linguistic conditions included semantically irreversible (“The boy is eating the apple”) and reversible (“The boy is pushing the girl”) sentences at three levels of syntactic complexity. As expected, patients performed well above chance on irreversible sentences, and at chance on reversible sentences of high complexity. Comprehension of reversible non-complex sentences ranged from nearly perfect to chance, and was highly correlated with offline measures of language comprehension. Lesion analysis revealed that comprehension deficits for reversible sentences were predicted by damage to the left temporal lobe. Although aphasic patients activated homologous areas in the right temporal lobe, such activation was not correlated with comprehension performance. Rather, patients with better comprehension exhibited increased activity in dorsal fronto-parietal regions. Correlations between performance and dorsal network activity occurred bilaterally during perception of sentences, and in the right hemisphere during a post-sentence memory delay. These results suggest that effortful reprocessing of perceived sentences in short-term memory can support improved comprehension in aphasia, and that strategic recruitment of alternative networks, rather than homologous takeover, may account for some findings of right hemisphere language activation in aphasia.
doi:10.1016/j.neuropsychologia.2013.03.007
PMCID: PMC3821997  PMID: 23566891
Language; Aphasia; Magnetoencephalography; Stroke; Syntax
15.  VISUAL ELEMENTS OF SUBJECTIVE PREFERENCE MODULATE AMYGDALA ACTIVATION 
Neuropsychologia  2007;45(10):2191-2200.
What are the basic visual cues that determine our preference towards mundane everyday objects? We previously showed that a highly potent cue is the nature of the object’s contour: people generally like objects with a curved contour compared with objects that have pointed features and a sharp-angled contour. This bias is hypothesized here to stem from an implicit perception of potential threat conveyed by sharp elements. Using human neuroimaging to test this hypothesis, we report that the amygdala, a brain structure that is involved in fear processing and has been shown to exhibit activation level that is proportional to arousal in general, is significantly more active for everyday sharp objects (e.g., a sofa with sharp corners) compared with their curved-contour counterparts. Therefore, our results indicate that a preference bias towards a visual object can be induced by low-level perceptual properties, independent of semantic meaning, via visual elements that on some level could be associated with threat. We further present behavioral results that provide initial support for the link between the sharpness of the contour and threat perception. Our brains might be organized to extract these basic contour elements rapidly for deriving an early warning signal in the presence of potential danger.
doi:10.1016/j.neuropsychologia.2007.03.008
PMCID: PMC4024389  PMID: 17462678
arousal; contour; fMRI; form; shape; threat
16.  Familiarity-based memory as an early cognitive marker of preclinical and prodromal AD 
Neuropsychologia  2013;51(6):1094-1102.
There is great interest in the development of cognitive markers that differentiate “normal” age-associated cognitive change from that of Alzheimer's disease (AD) in its prodromal (i.e., mild cognitive impairment; MCI) or even preclinical stages. Dual process models posit that recognition memory is supported by the dissociable processes of recollection and familiarity. Familiarity-based memory has generally been considered to be spared during normal aging, but it remains controversial whether this type of memory is impaired in early AD. Here, we describe findings of estimates of recollection and familiarity in young adults (YA), cognitively normal older adults (CN), and patients with amnestic-MCI (a-MCI). These measures in the CN and a-MCI patients were then related to a structural imaging biomarker of AD that has previously been demonstrated to be sensitive to preclinical and prodromal AD, the Cortical Signature of AD (ADsig). Consistent with much work in the literature, recollection, but not familiarity, was impaired in CN versus YA. Replicating our prior findings, a-MCI patients displayed impairment in both familiarity and recollection. Finally, the familiarity measure was correlated with the ADsig biomarker across the CN and a-MCI group, as well as within the CN adults alone. No other standard psychometric measure was as highly associated with the ADsig, suggesting that familiarity may be a sensitive biomarker of AD-specific brain changes in preclinical and prodromal AD and that it may offer a qualitatively distinct measure of early AD memory impairment relative to normal age-associated change.
doi:10.1016/j.neuropsychologia.2013.02.014
PMCID: PMC3742759  PMID: 23474075
Memory; Recollection; Familiarity; Alzheimer's disease; Medial temporal lobe; Preclinical Alzheimer's disease; Mild cognitive impairment
17.  Cognitive Reserve Moderates the Association Between Hippocampal Volume and Episodic Memory in Middle Age 
Neuropsychologia  2013;51(6):1124-1131.
Cognitive reserve is hypothesized to help people withstand greater brain pathology without manifesting clinical symptoms, and may be regarded as a preventive factor of dementia. It is unclear whether the effect of cognitive reserve is evident only among the older adults or after conversion to dementia, or if it can also be seen earlier in life before the prominent effects of cognitive aging become apparent. While finding a main effect of cognitive reserve on cognitive outcome may be consistent with the reserve hypothesis, in our view, it is unnecessary to invoke the idea of reserve if only a main effect is present. Rather, it is the interaction between a measure of reserve and a brain measure on cognitive outcome that is key for confirming that the effects of brain pathology affect people differently according to their cognitive reserve. We studied whether general cognitive ability at an average age of 20 years, as a direct measure of cognitive reserve, moderates the association between hippocampal volume and episodic memory performance in 494 middle-aged men ages 51 to 60. Whereas there was no statistically significant direct relationship between hippocampal volume and episodic memory performance in middle age, we found a statistically significant interaction such that there was a positive association between hippocampal volume and episodic memory only among people with lower general cognitive ability at age 20, i.e., lower levels of cognitive reserve. Our results provide support for the hypothesis that cognitive reserve moderates the relationship between brain structure and cognition in middle age, well before the onset of dementia.
doi:10.1016/j.neuropsychologia.2013.02.022
PMCID: PMC3660613  PMID: 23499725
cognitive reserve; general cognitive ability; episodic memory; hippocampus; verbal learning
18.  Unexpected novelty and familiarity orienting responses in lateral parietal cortex during recognition judgment 
Neuropsychologia  2013;51(6):1061-1076.
The role of lateral parietal cortex during recognition memory is heavily debated. We examined parietal activation during an Explicit Memory Cueing recognition paradigm that biases participants towards expecting novel or familiar stimuli on a trial-by-trial basis using anticipatory cues (“Likely Old”, “Likely New”), compared to trials with neutral cues (“????”). Three qualitatively distinct patterns were observed in the left lateral parietal cortex. An unexpected novelty response occurred in left anterior intraparietal cortex (IPS)/post-central gyrus (PoCG) in which greater activation was observed for new versus old materials following the “Likely Old” cue, but not following the “Likely New” cue. In contrast, anterior angular gyrus demonstrated an unexpected familiarity response with greater activation for old versus new materials following the “Likely New” cue, but not the “Likely Old” cue. Thus these two regions demonstrated increased responses that were selective for either new or old materials respectively, but only when they were unexpected. In contrast, a mid IPS area demonstrated greater response for whichever class of memoranda was unanticipated given the cue condition (an unexpected memory response). Analogous response patterns in regions outside of parietal cortex, and the results of a resting state connectivity analysis, suggested these three response patterns were associated with visuo-spatial orienting following unexpected novelty, source monitoring operations following unexpected familiarity, and general executive control processes following violated expectations. These findings support a Memory Orienting Model of the left lateral parietal cortex in which the region is linked to the investigation of unexpected novelty or familiarity in the environment.
doi:10.1016/j.neuropsychologia.2013.02.018
PMCID: PMC3690975  PMID: 23499719
episodic memory; parietal cortex; prefrontal cortex; decision biasing; external cues
19.  Subthalamic deep brain stimulation in Parkinson׳s disease has no significant effect on perceptual timing in the hundreds of milliseconds range 
Neuropsychologia  2014;57(100):29-37.
Bilateral, high-frequency stimulation of the basal ganglia (STN-DBS) is in widespread use for the treatment of the motor symptoms of Parkinson׳s disease (PD). We present here the first psychophysical investigation of the effect of STN-DBS upon perceptual timing in the hundreds of milliseconds range, with both duration-based (absolute) and beat-based (relative) tasks; 13 patients with PD were assessed with their STN-DBS ‘on’, ‘off’, and then ‘on’ again.
Paired parametric analyses revealed no statistically significant differences for any task according to DBS status. We demonstrate, from the examination of confidence intervals, that any functionally relevant effect of STN-DBS on relative perceptual timing is statistically unlikely. For absolute, duration-based timing, we demonstrate that the activation of STN-DBS may either worsen performance or have no effect, but that it is unlikely to lead to significant improvement.
Although these results are negative they have important implications for our understanding of perceptual timing and its relationship to motor functions within the timing network of the brain. They imply that the mechanisms involved in the perceptual processing of temporal information are likely to be functionally independent from those that underpin movement. Further, they suggest that the connections between STN and the subtantia nigra and globus pallidus are unlikely to be critical to beat-based perceptual timing.
Graphical abstract
Paired difference in performance thresholds according to DBS status. The first and third measures within each triplet show performance for that run with DBS ‘on’ compared to that with DBS ‘off’; values greater than zero denote poorer performance with DBS ‘on’, while those less than zero denote poorer performance with DBS ‘off’. The middle measure shows the difference in performance between the two runs with DBS ‘on’. Error bars denote confidence intervals.
Highlights
•We assessed the effect of STN-DBS on both duration- and beat-based perceptual timing.•DBS activation did not profoundly affect beat-based perceptual timing.•An effect of DBS on duration-based timing was not demonstrated but cannot be excluded.•Connections between STN and cerebellum are unlikely to be critical to perceptual timing.
doi:10.1016/j.neuropsychologia.2014.02.021
PMCID: PMC4022837  PMID: 24613477
Perceptual timing; Parkinson׳s disease; Subthalamic nucleus; Deep brain stimulation
20.  Phase locked neural activity in the human brainstem predicts preference for musical consonance 
Neuropsychologia  2014;58(100):23-32.
When musical notes are combined to make a chord, the closeness of fit of the combined spectrum to a single harmonic series (the ‘harmonicity’ of the chord) predicts the perceived consonance (how pleasant and stable the chord sounds; McDermott, Lehr, & Oxenham, 2010). The distinction between consonance and dissonance is central to Western musical form. Harmonicity is represented in the temporal firing patterns of populations of brainstem neurons. The current study investigates the role of brainstem temporal coding of harmonicity in the perception of consonance. Individual preference for consonant over dissonant chords was measured using a rating scale for pairs of simultaneous notes. In order to investigate the effects of cochlear interactions, notes were presented in two ways: both notes to both ears or each note to different ears. The electrophysiological frequency following response (FFR), reflecting sustained neural activity in the brainstem synchronised to the stimulus, was also measured. When both notes were presented to both ears the perceptual distinction between consonant and dissonant chords was stronger than when the notes were presented to different ears. In the condition in which both notes were presented to the both ears additional low-frequency components, corresponding to difference tones resulting from nonlinear cochlear processing, were observable in the FFR effectively enhancing the neural harmonicity of consonant chords but not dissonant chords. Suppressing the cochlear envelope component of the FFR also suppressed the additional frequency components. This suggests that, in the case of consonant chords, difference tones generated by interactions between notes in the cochlea enhance the perception of consonance. Furthermore, individuals with a greater distinction between consonant and dissonant chords in the FFR to individual harmonics had a stronger preference for consonant over dissonant chords. Overall, the results provide compelling evidence for the role of neural temporal coding in the perception of consonance, and suggest that the representation of harmonicity in phase locked neural firing drives the perception of consonance.
Highlights
•Harmonic structure of musical chords is encoded by phase locking in the brainstem.•Individual preference for consonance can be predicted by neural coding.•The neural coding of the harmonicity of a musical chord predicts how pleasant it is.•Monaural interactions generate neurally coded harmonics.•Generated harmonics enhance neural harmonicity and pleasantness of consonant chords.
doi:10.1016/j.neuropsychologia.2014.03.011
PMCID: PMC4040538  PMID: 24690415
Musical consonance; Individual differences; Auditory brainstem; Frequency following response; Pitch; Harmonicity
21.  Similar alterations in brain function for phonological and semantic processing to visual characters in Chinese dyslexia 
Neuropsychologia  2012;50(9):2224-2232.
Dyslexia in alphabetic languages has been extensively investigated and suggests a central deficit in orthography to phonology mapping in the left hemisphere. Compared to dyslexia in alphabetic languages, the central deficit for Chinese dyslexia is still unclear. Because of the logographic nature of Chinese characters, some have suggested that Chinese dyslexia should have larger deficits in the semantic system. To investigate this, Chinese children with reading disability (RD) were compared to typically developing (TD) children using functional magnetic resonance imaging (fMRI) on a rhyming judgment task and on a semantic association judgment task. RD children showed less activation for both tasks in right visual (BA18, 19) and left occipito-temporal cortex (BA 37), suggesting a deficit in visuo-orthographic processing. RD children also showed less activation for both tasks in left inferior frontal gyrus (BA44), which additionally showed significant correlations with activation of bilateral visuo-orthographic regions in the RD group, suggesting that the abnormalities in frontal cortex and in posterior visuo-orthographic regions may reflect a deficit in the connection between brain regions. Analyses failed to reveal larger differences between groups for the semantic compared to the rhyming task, suggesting that Chinese dyslexia is similarly impaired in the access to phonology and to semantics from the visual orthography.
doi:10.1016/j.neuropsychologia.2012.05.026
PMCID: PMC3987122  PMID: 22698991
Chinese dyslexia; Orthography; Phonology; Semantics
22.  Early gamma oscillations during rapid auditory processing in children with a language-learning impairment: Changes in neural mass activity after training 
Neuropsychologia  2013;51(5):990-1001.
Children with language-learning impairment (LLI) have consistently shown difficulty with tasks requiring precise, rapid auditory processing. Remediation based on neural plasticity assumes that the temporal precision of neural coding can be improved by intensive training protocols. Here, we examined the extent to which early oscillatory responses in auditory cortex change after audio-visual training, using combined source modeling and time-frequency analysis of the human electroencephalogram (EEG). Twenty-one elementary school students diagnosed with LLI underwent the intervention for an average of 32 days. Pre- and post-training assessments included standardized language/literacy tests and EEG recordings in response to fast-rate tone doublets. Twelve children with typical language development were also tested twice, with no intervention given. Behaviorally, improvements on measures of language were observed in the LLI group following completion of training. During the first EEG assessment, we found reduced amplitude and phase-locking of early (45–75 ms) oscillations in the gamma-band range (29–52 Hz), specifically in the LLI group, for the second stimulus of the tone doublet. Amplitude reduction for the second tone was no longer evident for the LLI children post-intervention, although these children still exhibited attenuated phase-locking. Our findings suggest that specific aspects of inefficient sensory cortical processing in LLI are ameliorated after training.
doi:10.1016/j.neuropsychologia.2013.01.011
PMCID: PMC3633611  PMID: 23352997
auditory temporal processing; computerized training; electroencephalography; phase locking; specific language impairment; spectral power
23.  Meaning first: A case for language-independent access to word meaning in the bilingual brain 
Neuropsychologia  2013;51(5):850-863.
This study aimed to determine how deeply a word is processed in the bilingual brain before the word’s language membership plays a role in lexical selection. In two ERP experiments, balanced Spanish-English bilinguals read lists of words and pseudowords in Spanish and English, and performed in each language 1) a language-specific lexical decision task, e.g., respond to real words in Spanish, and 2) a language-specific category decision tasks, e.g., respond to Spanish words that refer to a person. In Experiment 1, infrequent words elicited larger negativity between 350–650 msec post-stimulus onset for both target and non-target languages. This indicates that language membership did not block lexical access of non-target words, contrary to previous findings. In Experiment 2, we measured the onset of the target-category P300 as a way of determining if words from the non-target language were temporarily treated as targets. When Spanish was the target language, the ERP waveforms diverged early based on semantic category (people versus non-people), indicating that non-target ‘English people’ words were briefly treated as potential targets. This finding indicates that meaning was accessed prior to using language membership for lexical selection. However, when English was the target language, the waveforms diverged first based on language (Spanish versus English) then semantic category. We argue that the order in which meaning or language membership are accessed may be based on the frequency of use of a bilingual’s languages: the more frequently a language is used (English was more frequently used herein), the faster the words are identified as members of the language, and the greater interference it causes when it is not the target language. In brief, these findings make the case for a moment in processing when language membership matters less than meaning.
doi:10.1016/j.neuropsychologia.2013.01.017
PMCID: PMC3640818  PMID: 23376051
Bilingual lexical access; event-related potentials; language membership; lexical access; semantic categorisation; world frequency
24.  Getting it right: Word learning across the hemispheres 
Neuropsychologia  2013;51(5):825-837.
The brain is able to acquire information about an unknown word’s meaning from a highly constraining sentence context with minimal exposure. In this study, we investigate the potential contributions of the cerebral hemispheres to this ability. Undergraduates first read weakly or strongly constraining sentences completed by known or unknown (novel) words. Subsequently, their knowledge of these words was assessed via a lexical decision task in which they served as visual primes for lateralized target words varying in their semantic relationship to the primes (unrelated, identical or synonymous). As expected, smaller N400 amplitudes were seen for target words preceded by identical (vs. unrelated) known word primes, regardless of visual field of presentation. When Unknown words served as primes, N400 reductions to synonymous target words were observed only if the prime had appeared under High sentential constraint; targets appearing in the LVF/RH elicited a small N400 effect and modulation of a subsequent late positivity whereas those in the RVF/LH elicited modulation on the late positivity only. Unknown words initially seen in Low constraint contexts showed priming effects only in a late positivity and only in the RVF/LH. Strength of contextual constraint clearly seems to impact the hemispheres’ rapid acquisition of novel word meanings. N400 modulation for novel words under strong contextual constraint in the LVH/RH suggests that fast-mapped lexical representations may initially activate meanings that are weakly, distantly, associatively or thematically-related. More extensive and bilateral semantic processing seems to occur at longer processing latencies (post N400).
doi:10.1016/j.neuropsychologia.2013.01.027
PMCID: PMC3656665  PMID: 23416731
ERPs; N400; word learning; fast-mapping; cerebral hemispheres; language; priming
25.  Neural competition as a developmental process: Early hemispheric specialization for word processing delays specialization for face processing 
Neuropsychologia  2013;51(5):950-959.
Little is known about the impact of learning to read on early neural development for word processing and its collateral effects on neural development in non-word domains. Here, we examined the effect of early exposure to reading on neural responses to both word and face processing in preschool children with the use of the Event Related Potential (ERP) methodology. We specifically linked children’s reading experience (indexed by their sight vocabulary) to two major neural markers: the amplitude differences between the left and right N170 on the bilateral posterior scalp sites and the hemispheric spectrum power differences in the γ band on the same scalp sites. The results showed that the left-lateralization of both the word N170 and the spectrum power in the γ band were significantly positively related to vocabulary. In contrast, vocabulary and the word left-lateralization both had a strong negative direct effect on the face right-lateralization. Also, vocabulary negatively correlated with the right-lateralized face spectrum power in the γ band even after the effects of age and the word spectrum power were partialled out. The present study provides direct evidence regarding the role of reading experience in the neural specialization of word and face processing above and beyond the effect of maturation. The present findings taken together suggest that the neural development of visual word processing competes with that of face processing before the process of neural specialization has been consolidated.
doi:10.1016/j.neuropsychologia.2013.02.006
PMCID: PMC3756286  PMID: 23462239
Neural development; Face processing; Visual word processing; ERP

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