Dissociating explicit and implicit category knowledge with fMRI

Neuroimaging of healthy volunteers identified separate neural systems supporting the expression of category knowledge depending on whether the learning mode was intentional or incidental. The same visual category was learned either intentionally or implicitly by two separate groupsof participants. During a categorization test, functional magnetic resonance imaging (fMRI) was used to compare brain activity evoked by category members and nonmembers. After implicit learning, when participants had learned the category incidentally, decreased occipital activity was observed for novel categorical stimuli compared with noncategorical stimuli. In contrast, after intentional learning, novel categorical stimuli evoked increased activity in the hippocampus, right prefrontal cortex, left inferior temporal cortex, precuneus, and posterior cingulate. Even though the categorization test was identical in the two conditions, the differences in brain activity indicate differing representations of category knowledge depending on whether the category had been learned intentionally or implicitly.     

The emotional paradox: Dissociation between explicit and implicit processing of emotional prosody in schizophrenia

People with schizophrenia show well-replicated deficits on tasks of explicit recognition of emotional prosody. However it remains unclear whether they are still sensitive to the implicit cues of emotional prosody, particularly when they exhibit high levels of social anhedonia. A dual processing model suggesting a dissociation between the neural networks involved in explicit and implicit recognition of emotional prosody has yet to be validated.21 participants with schizophrenia and 21 controls were recruited. In the explicit recognition task, individuals listened to semantically neutral words pronounced with two different emotions and judged their emotional prosody. In the vocal emotional Stroop task, patients and controls listened to words with a positive or negative emotional valence pronounced with congruent or incongruent emotional prosody and judged their emotional content. Patients were also assessed with the Chapman Anhedonia Questionnaire and the Schizophrenic Communication Disorders scale.Individuals with schizophrenia were impaired in their explicit recognition of emotional prosody related to controls. In contrast, they showed a vocal emotional Stroop effect that was identical to controls for reaction time and greater for accuracy: patients were still sensitive to implicit emotional prosody. In addition the vocal emotional Stroop score increased with social anhedonia but was unrelated to communication disorders.Whereas explicit vocal affect recognition is impaired, implicit processing of emotional prosody seems to be preserved in schizophrenia. Our results provide evidence that at a behavioural level, the implicit and explicit processing of emotional prosody can be dissociated. Remediation of emotional prosody recognition in schizophrenia should target cognitive rather than sensory processes.     

Evidence of developmental differences in implicit sequence learning: an fMRI study of children and adults

Prevailing theories of implicit or unaware learning propose a developmental invariance model, with implicit function maturing early in infancy or childhood despite prolonged improvements in explicit or intentional learning and memory systems across childhood. Neuroimaging studies of adult visuomotor sequence learning have associated fronto-striatal brain regions with implicit learning of spatial sequences. Given evidence of continued development in these brain regions during childhood, we compare implicit sequence learning in adults and 7- to 11-year-old children to examine potential developmental differences in the recruitment of fronto-striatal circuitry during implicit learning. Participants performed a standard serial reaction time task. Stimuli alternately followed a fixed 10-step sequence of locations or were presented in a pseudorandom order of locations. Adults outperformed children, achieving a significantly larger sequence learning effect and showing learning more quickly than children. Age-related differences in activity were observed in the premotor cortex, putamen, hippocampus, inferotemporal cortex, and parietal cortex. We observed differential recruitment of cortical and subcortical motor systems between groups, presumably reflecting age differences in motor response execution. Adults showed greater hippocampal activity for sequence trials, whereas children demonstrated greater signal during random trials. Activity in the right caudate correlated significantly with behavioral measures of implicit learning for both age groups, although adults showed greater signal change than children overall, as would be expected given developmental differences in sequence learning magnitude. These results challenge the idea of developmental invariance in implicit learning and instead support a view of parallel developments in implicit and explicit learning systems.     

抑郁症患者外显性与内隐性情绪处理的脑功能磁共振研究

目的以健康者为对照,利用脑功能磁共振研究抑郁症患者的外显性和内隐性情绪处理过程。方法2006年12月-2007年12月,收集临床诊断抑郁症患者14例（DSM-Ⅳ标准）。14例健康志愿者作为对照。采用传统的组块设计,采集患者在高兴与悲伤2组脸像刺激下,判断表情的外显性情绪处理和判断性别的内隐性情绪处理过程的脑功能磁共振图像,利用SPM2统计软件计算出个体及组内在不同表情刺激和不同任务操作下激活的脑功能区。结果①抑郁症患者判断表情时,悲伤脸像的主要激活区位于顶叶、海马旁回、基底节、梭状回、丘脑、岛叶、前扣带回及胼胝体下回;高兴脸像激活区仅有前扣带回;②抑郁症患者判断性别任务时,悲伤脸像激活区分布在额中回、顶下小叶、前扣带回、颞中回;高兴脸像未见明显脑功能激活区。结论①抑郁症患者外显性与内隐性情绪刺激的脑内加工过程不同,悲伤脸像脑功能区激活均强于高兴脸像;②与健康人群相比,不同情绪处理过程中,抑郁症患者的高兴脸像激活区较弱,而悲伤脸像的激活明显增强。提示抑郁症患者情绪处理的神经系统存在异常,对正性情绪的神经反应减弱,而对负性情绪的神经反应增强。     

Mismatch negativity (MMN) stands at the crossroads between explicit and implicit emotional processing

The amygdala is known as a key brain region involved in the explicit and implicit processing of emotional faces, and plays a crucial role in salience detection. Not until recently was the mismatch negativity (MMN), a component of the event‐related potentials to an odd stimulus in a sequence of stimuli, utilized as an index of preattentive salience detection of emotional voice processing. However, their relationship remains to be delineated. This study combined the fMRI scanning and event‐related potential recording by examining amygdala reactivity in response to explicit and implicit (backward masked) perception of fearful and angry faces, along with recording MMN in response to the fearfully and angrily spoken syllables dada in healthy subjects who varied in trait anxiety (STAI‐T). Results indicated that the amplitudes of fearful MMN were positively correlated with left amygdala reactivity to explicit perception of fear, but negatively correlated with right amygdala reactivity to implicit perception of fear. The fearful MMN predicted STAI‐T along with left amygdala reactivity to explicit fear, whereas the association between fearful MMN and STAI‐T was mediated by right amygdala reactivity to implicit fear. These findings suggest that amygdala reactivity in response to explicit and implicit threatening faces exhibits opposite associations with emotional MMN. In terms of emotional processing, MMN not only reflects preattentive saliency detection but also stands at the crossroads of explicit and implicit perception. Hum Brain Mapp 38:140–150, 2017. © 2016 Wiley Periodicals, Inc.     

Cerebral hemispheric differences in memory of emotional and non-emotional words in normal individuals

The present study was designed to examine the cerebral hemispheric differences in memory of positive, negative and non-emotional words using a new method of successive presentation to each visual half-field in which perception of each item was nearly perfect thereby allowing laterality differences for effects of emotion on memory to emerge unconfounded by perception (Experiment 1). This procedure was compared with traditional perceptual identification (Experiment 2). In Experiment 1, 12 words were presented successively in each half field in each trial followed by free recall at the end of the trial. The results showed that recall of positive and negative emotional words was better than that of non-emotional words in both visual fields. Recall of positive and negative emotional words was not different in left and right visual fields (RVFs) although the recall of non-emotional words was better in the RVF than in the left visual field (LVF). The differences in recall between emotional and non-emotional words was greater in the LVF than in the RVF. Experiment 2 used the more traditional method of perceptual identification following each visual half-field presentation of a single item. Perceptual identification was better in the RVF than the LVF in each word condition. There were no visual field differences in perceptual identification between emotional and non-emotional words, as there was for memory in Experiment 1. The results supported the hypothesis that explicit memory for emotional words was dependent more on the right hemisphere, whereas perception of both emotional and non-emotional words was more dependent on the left hemisphere.     

Brain responses of explicit and implicit memory: An event-related potential study

Implicit memory is acquired by an unintentional or unconscious learning. Recognition memory involves either automatic knowing or consciously controlled remembering. We provided an event-related potential paradigm capable of differentiating memory for the explicitly learned, implicitly learned and unstudied materials. In the explicit memory, we obtained both frontal (controlled retrieval) and parietal (recollection) old/new effects. In the implicit memory, we found persistent occipitotemporal activation (visual priming) and late attenuation in the temporoparietooccipital (repetition suppression). Event-related potential provides an insight into the dissociable mechanism of memory function that supports the dual process model with an enhanced temporal resolution on the dynamic process of both explicit perceptual learning and implicit perceptual priming.     

Levels of emotional awareness and autism: An fMRI study

Abstract

Autism is associated with an inability to identify and distinguish one's own feelings. We assessed this inability using alexithymia and empathy questionnaires, and used fMRI to investigate brain activity while introspecting on emotion. Individuals with high functioning autism/Asperger syndrome (HFA/AS) were compared with matched controls. Participants rated stimuli from the International Affective Picture System twice, once according to the degree of un/pleasantness that the pictures induced, and once according to their color balance. The groups differed significantly on both alexithymia and empathy questionnaires. Alexithymia and lack of empathy were correlated, indicating a link between understanding one's own and others’ emotions. For both groups a strong relationship between questionnaire scores and brain activity was found in the anterior insula (AI), when participants were required to assess their feelings to unpleasant pictures. Regardless of self-reported degree of emotional awareness, individuals with HFA/AS differed from controls when required to introspect on their feelings by showing reduced activation in self-reflection/mentalizing regions. Thus, we conclude that difficulties in emotional awareness are related to hypoactivity in AI in both individuals with HFA/AS and controls, and that the particular difficulties in emotional awareness in individuals with HFA/AS are not related to their impairments in self-reflection/mentalizing.     

Levels of emotional awareness and autism: an fMRI study

Autism is associated with an inability to identify and distinguish one's own feelings. We assessed this inability using alexithymia and empathy questionnaires, and used fMRI to investigate brain activity while introspecting on emotion. Individuals with high functioning autism/Asperger syndrome (HFA/AS) were compared with matched controls. Participants rated stimuli from the International Affective Picture System twice, once according to the degree of un/pleasantness that the pictures induced, and once according to their color balance. The groups differed significantly on both alexithymia and empathy questionnaires. Alexithymia and lack of empathy were correlated, indicating a link between understanding one's own and others' emotions. For both groups a strong relationship between questionnaire scores and brain activity was found in the anterior insula (AI), when participants were required to assess their feelings to unpleasant pictures. Regardless of self-reported degree of emotional awareness, individuals with HFA/AS differed from controls when required to introspect on their feelings by showing reduced activation in self-reflection/mentalizing regions. Thus, we conclude that difficulties in emotional awareness are related to hypoactivity in AI in both individuals with HFA/AS and controls, and that the particular difficulties in emotional awareness in individuals with HFA/AS are not related to their impairments in self-reflection/mentalizing.     

Cerebral asymmetries in monozygotic twins: An fMRI study

Lateralization for language, spatial judgment and face processing was assessed in 42 pairs of identical twins, 21 discordant and 21 concordant pairs for handedness, using fMRI. Individual laterality indices were calculated based on the observed activation patterns. All tasks showed expected asymmetry, favoring the left-hemisphere for language and the right-hemisphere for spatial judgment and face processing. The intra-class correlations on the laterality indices were significant only on the language task and only for the concordant group, but not the discordant group, suggesting a stronger genetic influence for language asymmetry in concordant twins. The expected asymmetry was greater for the concordant group only on the language task. The difference was not significant, but conformed quite well to Annett's genetic model, which assumes a right-shift (RS) gene with one allele (RS+) biasing toward right-handedness and left-cerebral language dominance, and the other (RS−) leaving both asymmetries to chance. The model also assumes that the genetic influence is additive for handedness but dominant-recessive for left-cerebral language dominance, which explains the high concordance for language dominance in twins discordant for handedness. Our data suggest that the same gene has no influence on right-hemisphere dominance for spatial judgment or face processing, and offer little support for mirror-imaging in MZ twins other than that due to chance.     

Attentional modulation of emotional stimulus processing: an fMRI study using emotional expectancy

We used emotional expectancy to study attentional modulation in the processing of emotional stimuli. During functional magnetic resonance imaging (fMRI), volunteers saw emotional and neutral expectancy cues signaling the subsequent presentation of corresponding emotional or neutral pictorial stimuli. As a control, emotional and neutral pictures were presented without preceding expectancy cue, resulting in a 2 x 2 factorial design with the factors "expectancy" and "emotion." Statistical analysis revealed a significant positive interaction effect between these factors in the medial prefrontal cortex (MPFC, Brodmann area [BA] 9/10), amygdala, and dorsal midbrain. In all these regions, expectancy augmented the neural response to emotional but not to neutral pictures. Time course analysis of raw data suggests that this augmented activation was not preceded by baseline increases in MPFC and amygdala during the period of emotional expectancy. In a post-scanning session, the paradigm was presented for a second time to allow emotional intensity rating. Again, a significant interaction between expectancy and emotion was observed, with intensity ratings specifically enhanced in emotional photographs preceded by expectancy. There was a positive correlation between intensity ratings and blood oxygenation level-dependent (BOLD) signals in the left amygdala. We conclude that specific components of the emotion network show enhanced activation in response to emotional stimuli when these are preceded by expectancy. This enhancement effect is not present in neutral pictures and might parallel accentuated subjective feeling states.     

Reduced implicit and explicit sequence learning in first-episode schizophrenia

A high prevalence of deficits in explicit learning has been reported for schizophrenic patients, but it is less clear whether these patients are impaired in implicit learning. Deficits in implicit learning indicative of a fronto-striatal dysfunction have been reported using a serial reaction-time task (SRT), but the impact of typical neuroleptic medication and chronicity remains controversial. The present study compared 37 patients with first-episode schizophrenia treated with atypical neuroleptics and 37 healthy matched control participants on two sequence learning tasks: a modified SRT for implicit sequence learning and a serial generation task (SGT) for explicit sequence learning. The two tasks were designed to be procedurally equivalent, in order to provide better comparability between implicit and explicit performance. Although unaffected in global cognitive functioning, schizophrenic patients were significantly impaired in implicit and explicit sequence learning. Deficient sequence learning in schizophrenic patients was neither related to psychopathology nor to chlorpromazine equivalent daily dosage. As performance was impaired even though patients were exclusively treated with atypical neuroleptics, the present findings concur with converging evidence of a sequence learning deficit inherent in schizophrenia. This deficit would be consistent with a fronto-striatal dysfunction and might constitute a crucial factor for the acquisition of new information.     

Right hemisphere dysfunction and emotional processing in ALS: an fMRI study

Emotional processing may be abnormal in amyotrophic lateral sclerosis (ALS). Our aim was to explore functional anatomical correlates in the processing of aversive information in ALS patients. We examined the performance of nine non-demented ALS patients and 10 healthy controls on two functional MRI (fMRI) tasks, consisting of an emotional attribution task and a memory recognition task of unpleasant versus neutral stimuli. During the emotional decision task, subjects were asked to select one of three unpleasant or neutral words. During the memory task, subjects were asked to recognize words presented during the previous task. Controls showed, as expected, greater activation in the right middle frontal gyrus during selection of unpleasant than neutral words, and a greater activation mainly in right-sided cerebral areas during the emotional recognition task. Conversely, patients showed a general increase in activation of the left hemisphere, and reduced activation in right hemisphere in both emotional tasks. Such findings may suggest extra-motor neurodegeneration involving key circuits of emotions, mostly negative, commonly involved in FTD.     

A study of parallel implicit and explicit information processing in patients with obsessive-compulsive disorder

OBJECTIVE: This study examined implicit sequence learning in patients with obsessive-compulsive disorder (OCD) under dual-task conditions. Frontal-striatal networks support implicit learning and are implicated in the pathophysiology of OCD. Neuroimaging data suggest that during implicit learning, OCD patients use neural systems normally active during explicit learning to compensate for striatal dysfunction. METHOD: The authors examined implicit sequence learning in 25 OCD patients and 25 healthy comparison subjects using a dual-task paradigm, with subjects simultaneously engaged in an explicit memory task and an implicit learning task. They predicted that implicit learning in OCD subjects would be disrupted because concurrent explicit information-processing demands would prevent use of compensatory processes. RESULTS: OCD patients failed to show evidence of implicit learning and exhibited a significant deficit in comparison with healthy subjects. CONCLUSIONS: These results are consistent with the hypothesis that concurrent explicit and implicit information-processing demands interfere with implicit learning in OCD patients.     

Objects automatically potentiate action: an fMRI study of implicit processing

Behavioural data have shown that the perception of an object automatically potentiates motor components (affordances) of possible actions toward that object, irrespective of the subject's intention. We carried out an event-related fMRI study to investigate the influence of the intrinsic properties of an object on motor responses which were either compatible or incompatible with the action that the object affords. The subjects performed power or precision grip responses based on the categorization of objects into natural or man-made. The objects were either 'small' (usually grasped with a precision grip) or 'large' (usually grasped with a power grip). As expected, the motor responses were fastest to objects that afforded the same grip (congruent) and slowest to objects that afforded the other grip (incongruent). Imaging revealed activations which covaried with compatibility in the parietal, dorsal premotor and inferior frontal cortex. We suggest that the greater the difference in reaction times between congruent and incongruent trials, the greater the competition between the action afforded by the object and the action specified by the task, and thus the greater the activation within this network.     

Implicit and explicit emotional processing in Parkinson's disease

Recent semantic priming investigations conducted by Spicer, Brown and Gorell (1994) and McDonald, Brown and Gorell (1996) reported hyperpriming in persons with Parkinson's disease (PD) and provided evidence supporting impaired set-shifting as the underlying cause. This paper discusses notable priming behaviours exhibited by the normal control groups employed by Spicer and McDonald and colleagues. The argument is developed that these studies only measured attention-dependent semantic processing and, therefore, their results do not speak to issues of automatic semantic activation in PD as originally proposed by Spicer et al. nor to putative deficits in set-shifting as proposed by McDonald et al. The implications for future priming research in PD are also discussed.     

Age-related differences in the medial temporal lobe responses to emotional faces as revealed by fMRI

Age-related differences involved in the neural substrates of emotional face perception were investigated in young and old healthy volunteers. The subjects were scanned using functional magnetic resonance imaging while they were judging the gender of faces with negative, positive, or neutral emotional valence. The results showed that both the predominant activation in young subjects and reduced activity in old subjects contributed to a significant age difference in the left amygdala during the perception of negative faces. Activity in the right parahippocampal gyrus during the perception of positive faces diminished with advancing age. Neural activity in the angular gyrus and lingual gyrus of the right hemisphere was reduced in the old subjects during the perception of positive faces. There was no region where old subjects had greater activity than young subjects during the task. In old subjects, the overall activity in the right hippocampus during the task correlated negatively with age, whereas the activity in the right parahippocampal gyrus correlated positively with neuropsychological performance. There was no significant correlation between subjects' characteristics and signal change in young subjects. These results indicate the age-associated vulnerability of the medial temporal lobe structures including the amygdala, hippocampus, and parahippocampal gyrus during face perception. The dissociation with reduced activity in the left amygdala and the right parahippocampal gyrus may suggest that aging differentially affects neural responses to faces with negative or positive emotional valence. The parieto-occipital lobe, which has been found to be involved in face processing, also showed a functional decline associated with aging.     

Error and deviance processing in implicit and explicit sequence learning

In this experiment, we examined the extent to which error-driven learning may operate under implicit learning conditions. We compared error monitoring in a sequence learning task in which stimuli consisted of regular, irregular, or random sequences. Subjects were either informed (explicit condition) or not informed (implicit condition) about the existence of the sequence. For both conditions, reaction times were faster to stimuli from regular sequences than from random sequences, thus supporting the view that sequence learning occurs irrespective of learning condition. Response-locked event-related potentials (ERPs) showed a pronounced ERN/Ne, thereby signaling the detection of committed errors. Deviant stimuli from irregular sequences elicited an N2b component that developed in the course of the experiment, albeit faster for explicit than implicit learners. This observation supports the view that deviant events acquire the status of perceived errors during explicit and implicit learning, and thus, an N2b is generated resembling the ERN/Ne to committed errors. While performing the task, expectations about upcoming events are generated, compared to the actual events, and evaluated on the dimension "better or worse than expected." The accuracy of this process improves with learning, as shown by a gradual increase in N2b amplitude as a function of learning. Additionally, a P3b, which is thought to mirror conscious processing of deviant stimuli and is related to updating of working memory representations, was found for explicit learners only.     

The emotional modulation of cognitive processing: an fMRI study

The functional neuroanatomy of visual processing of surface features of emotionally valenced pictorial stimuli was examined in normal human subjects using functional magnetic resonance imaging (fMRI). Pictorial stimuli were of two types: emotionally negative and neutral pictures. Task performance was slower for the negatively valenced than for the neutral pictures. Significant blood oxygen level dependent (BOLD) increases occurred in the medial and dorsolateral prefrontal cortex, midbrain, substantia innominata, and/or amygdala, and in the posterior cortical visual areas for both stimulus types. Increases were greater for the negatively valenced stimuli. While there was a small but significant BOLD decrease in the subgenual prefrontal cortex, which was larger in response to the negatively valenced pictures, there was an almost complete absence of other decreases prominently seen during the performance of demanding cognitive tasks [Shulman, G. L., Fiez, J. A., Corbetta, M., Buckner, R. L., Miezin, F. M., Raichle, M. E., & Petersen, S. E. (1997). Common blood flow changes across visual tasks: II. Decreases in cerebral cortex. Journal of Cognitive Neuroscience, 9, 648--663]. These results provide evidence that the emotional valence and arousing nature of stimuli used during the performance of an attention-demanding cognitive task are reflected in discernable, quantitative changes in the functional anatomy associated with task performance.