Differential amygdala activation during emotional decision and recognition memory tasks using unpleasant words: an fMRI study

This study used fMRI to examine the response of the amygdala in the evaluation and short-term recognition memory of unpleasant vs. neutral words in nine right-handed healthy adult women. To establish specificity of the amygdala response, we examined the fMRI BOLD signal in one control region (visual cortex). Alternating blocks of unpleasant and neutral trials were presented. During the emotional decision task, subjects viewed sets of three unpleasant or three neutral words while selecting the most unpleasant or neutral word, respectively. During the memory task, subjects identified words that were presented during the emotional decision task (0.50 probability). Images were detrended, filtered, and coregistered to standard brain coordinates. The Talairach coordinates for the center of the amygdala were chosen before analysis. The BOLD signal at this location in the right hemisphere revealed a greater amplitude signal for the unpleasant relative to the neutral words during the emotional decision but not the memory task, confirmed by Time Course x Word Condition ANOVAs. These results are consistent with the memory modulatory view of amygdala function, which suggests that the amygdala facilitates long-term, but not short-term, memory consolidation of emotionally significant material. The control area showed only an effect for Time Course for both the emotional decision and memory tasks, indicating the specificity of the amygdala response to the evaluation of unpleasant words. Moreover, the right-sided amygdala activation during the unpleasant word condition was strongly correlated with the BOLD response in the occipital cortex. These findings corroborate those by other researchers that the amygdala can modulate early processing of visual information in the occipital cortex. Finally, an increase in subject's state anxiety (evaluated by questionnaire) while in the scanner correlated with amygdala activation under some conditions.     

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.     

The influence of positive and negative emotional associations on semantic processing in depression: An fMRI study

In depression, patients suffer from emotional and cognitive deficits, among others in semantic processing. If these semantic deficits are cognitive or interact with emotional dysfunctions, is still an open question. The aim of the current study was to investigate the influence of emotional valence on the neural correlates of semantic priming in major depression. In a lexical decision task, positive, negative, and neutral word pairs were presented during fMRI measurement. Nineteen inpatients and 19 demographically matched controls were recruited. Behaviorally, positive and neutral valence induced a priming effect whereas negative valence induced no effect (controls) or even inhibition (slower RT for related stimuli) in patients. At the neural level, the semantic relation effect revealed similar neural activation in right middle frontal regions for patients and controls. Group differences emerged in the right fusiform gyrus and the ACC. Activity associated with positive valence differed at the DLPFC and amygdala and for negative valence at putamen and cerebellum. The activation of amygdala and DLPFC correlated negatively with the severity of depression. To conclude, semantic processing deficits in depression are modulated by emotional valence of the stimulus on the behavioral as well as on neural level in right‐lateralized prefrontal areas and the amygdala. The results highlighted an influence of depression severity on emotion information processing as the severity of symptoms correlated negatively with neural responses to positively and negatively valenced information. Hence, the dysfunctional emotion processing may further enhance the cognitive deficits in depression. Hum Brain Mapp 35:471–482, 2014. © 2012 Wiley Periodicals, Inc.     

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.     

Imagined rotations of self versus objects: an fMRI study

This study used functional magnetic resonance imaging (fMRI) to investigate the neural mechanisms underlying two types of spatial transformations: imagined object rotations and imagined rotations of the self about an object. Participants viewed depictions of single three-dimensional Shepard--Metzler objects situated within a sphere. A T-shaped prompt appeared outside of the sphere at different locations across trials. In the object rotation task, participants imagined rotating the object so that one of its ends was aligned with the prompt. They then judged whether a textured portion of the object would be visible in its new orientation. In the self rotation task, they imagined rotating themselves to the location of the T-prompt, and then judged whether a textured portion of the object would be visible from the new viewpoint. Activation in both tasks was compared to respective control conditions in which identical judgments were made without rotation. A direct comparison of self and object rotation tasks revealed activation spreading from left premotor to left primary motor (M1) cortex (areas 6/4) for imagined object rotations, but not imagined self rotations. In contrast, the self rotation task activated left supplementary motor area (SMA; area 6). In both transformations, activation also occurred in other regions. These findings provide evidence for multiple spatial-transformation mechanisms within the human cognitive system.     

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.     

Differences in auditory processing of words and pseudowords: an fMRI study

Although there has been great interest in the neuroanatomical basis of reading, little attention has been focused on auditory language processing. The purpose of this study was to examine the differential neuroanatomical response to the auditory processing of real words and pseudowords. Eight healthy right-handed participants performed two phoneme monitoring tasks (one with real word stimuli and one with pseudowords) during a functional magnetic resonance imaging (fMRI) scan with a 4.1 T system. Both tasks activated the inferior frontal gyrus (IFG), the posterior superior temporal gyrus (pSTG) and the inferior parietal lobe (IPL). Pseudoword processing elicited significantly more activation within the posterior cortical regions compared with real word processing. Previous reading studies have suggested that this increase is due to an increased demand on the lexical access system. The left inferior frontal gyrus, on the other hand, did not reveal a significant difference in the amount of activation as a function of stimulus type. The lack of a differential response in IFG for auditory processing supports its hypothesized involvement in grapheme to phoneme conversion processes. These results are consistent with those from previous neuroimaging reading studies and emphasize the utility of examining both input modalities (e.g., visual or auditory) to compose a more complete picture of the language network.     

The self across the senses: an fMRI study of self-face and self-voice recognition

There is evidence that the right hemisphere is involved in processing self-related stimuli. Previous brain imaging research has found a network of right-lateralized brain regions that preferentially respond to seeing one''s own face rather than a familiar other. Given that the self is an abstract multimodal concept, we tested whether these brain regions would also discriminate the sound of one''s own voice compared to a friend''s voice. Participants were shown photographs of their own face and friend''s face, and also listened to recordings of their own voice and a friend''s voice during fMRI scanning. Consistent with previous studies, seeing one''s own face activated regions in the inferior frontal gyrus (IFG), inferior parietal lobe and inferior occipital cortex in the right hemisphere. In addition, listening to one''s voice also showed increased activity in the right IFG. These data suggest that the right IFG is concerned with processing self-related stimuli across multiple sensory modalities and that it may contribute to an abstract self-representation.     

Brain activity during expectancy of emotional stimuli: an fMRI study

We studied the neural activation associated with the expectancy of emotional stimuli using whole brain fMRI. Fifteen healthy subjects underwent fMRI scanning during which they performed a warned reaction task using emotional pictures carrying pleasant, unpleasant, or neutral content. The task involved an expected or unexpected condition. Data were analyzed by comparing the images acquired under the different conditions. In the expected condition, compared with the unexpected condition, significant activation was observed in the medial, inferior and dorsolateral prefrontal cortex. Whereas the expectancy of pleasant stimuli produced activation in the left dorsolateral and left medial prefrontal cortex as well as in the right cerebellum, the expectancy of unpleasant stimuli produced activation in the right inferior and right medial prefrontal cortex, the right amygdala, the left anterior cingulate cortex, and bilaterally in the visual cortex. These results suggest that the expectancy of emotional stimuli is mediated by the prefrontal area including the medial, inferior, and dorsolateral prefrontal cortex. Furthermore, our data suggest that left frontal activation is associated with the expectancy of pleasant stimuli and that right frontal activation is associated with the expectancy of unpleasant stimuli. Finally, our findings suggest that the amygdala and anterior cingulate cortex may play an important role in the expectancy of unpleasant stimuli and that the input of this negative information is modulated by these specific brain areas.     

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.     

Integration of cross-modal emotional information in the human brain: an fMRI study

The interaction of information derived from the voice and facial expression of a speaker contributes to the interpretation of the emotional state of the speaker and to the formation of inferences about information that may have been merely implied in the verbal communication. Therefore, we investigated the brain processes responsible for the integration of emotional information originating from different sources. Although several studies have reported possible sites for integration, further investigation using a neutral emotional condition is required to locate emotion-specific networks. Using functional magnetic resonance imaging (fMRI), we explored the brain regions involved in the integration of emotional information from different modalities in comparison to those involved in integrating emotionally neutral information. There was significant activation in the superior temporal gyrus (STG); inferior frontal gyrus (IFG); and parahippocampal gyrus, including the amygdala, under the bimodal versus the unimodal condition, irrespective of the emotional content. We confirmed the results of previous studies by finding that the bimodal emotional condition elicited strong activation in the left middle temporal gyrus (MTG), and we extended this finding to locate the effects of emotional factors by using a neutral condition in the experimental design. We found anger-specific activation in the posterior cingulate, fusiform gyrus, and cerebellum, whereas we found happiness-specific activation in the MTG, parahippocampal gyrus, hippocampus, claustrum, inferior parietal lobule, cuneus, middle frontal gyrus (MFG), IFG, and anterior cingulate. These emotion-specific activations suggest that each emotion uses a separate network to integrate bimodal information and shares a common network for cross-modal integration.     

Switching between colors and shapes on the basis of positive and negative feedback: an fMRI and EEG study on feedback-based learning

A crucial element of testing hypotheses about rules for behavior is the use of performance feedback. In this study, we used fMRI and EEG to test the role of medial prefrontal cortex (PFC) and dorsolateral (DL) PFC in hypothesis testing using a modified intradimensional/extradimensional rule shift task. Eighteen adults were asked to infer rules about color or shape on the basis of positive and negative feedback in sets of two trials. Half of the trials involved color-to-color or shape-to-shape trials (intradimensional switches; ID) and the other half involved color-to-shape or shape-to-color trials (extradimensional switches; ED). Participants performed the task in separate fMRI and EEG sessions. ED trials were associated with reduced accuracy relative to ID trials. In addition, accuracy was reduced and response latencies increased following negative relative to positive feedback. Negative feedback resulted in increased activation in medial PFC and DLPFC, but more so for ED than ID shifts. Reduced accuracy following negative feedback correlated with increased activation in DLPFC, and increased response latencies following negative feedback correlated with increased activation in medial PFC. Additionally, around 250 msec following negative performance feedback participants showed a feedback-related negative scalp potential, but this potential did not differ between ID and ED shifts. These results indicate that both medial PFC and DLPFC signal the need for performance adjustment, and both regions are sensitive to the increased demands of set shifting in hypothesis testing.     

The influence of self-awareness on emotional memory formation: an fMRI study

Evidence from functional neuroimaging studies of emotional perception shows that when attention is focused on external features of emotional stimuli (external perceptual orienting—EPO), the amygdala is primarily engaged, but when attention is turned inwards towards one’s own emotional state (interoceptive self-orienting—ISO), regions of the salience network, such as the anterior insula (AI) and the dorsal anterior cingulate cortex (dACC), also play a major role. Yet, it is unknown if ISO boosts the contributions of AI and dACC not only to emotional ‘perception’ but also to emotional ‘memory’. To investigate this issue, participants were scanned with functional magnetic resonance imaging (fMRI) while viewing emotional and neutral pictures under ISO or EPO, and memory was tested several days later. The study yielded three main findings: (i) emotion boosted perception-related activity in the amygdala during both ISO and EPO and in the right AI exclusively during ISO; (ii) emotion augmented activity predicting subsequent memory in AI and dACC during ISO but not during EPO and (iii) high confidence memory was associated with increased amygdala–dACC connectivity, selectively for ISO encoding. These findings show, for the first time, that ISO promotes emotional memory formation via regions associated with interoceptive awareness of emotional experience, such as AI and dACC.     

Neural correlates of semantic priming for ambiguous words: an event-related fMRI study

We investigated the neural correlates of semantic priming by using event-related fMRI to record blood oxygen level dependent (BOLD) responses while participants performed speeded lexical decisions (word/nonword) on visually presented related versus unrelated prime-target pairs. A long stimulus onset asynchrony of 1000 ms was employed, which allowed for increased controlled processing and selective frequency-based ambiguity priming. Conditions included an ambiguous word prime (e.g. bank) and a target related to its dominant (e.g. money) or subordinate meaning (e.g. river). Compared to an unrelated condition, primed dominant targets were associated with increased activity in the LIFG, the right anterior cingulate and superior temporal gyrus, suggesting postlexical semantic integrative mechanisms, while increased right supramarginal activity for the unrelated condition was consistent with expectancy based priming. Subordinate targets were not primed and were associated with reduced activity primarily in occipitotemporal regions associated with word recognition, which may be consistent with frequency-based meaning suppression. These findings provide new insights into the neural substrates of semantic priming and the functional-anatomic correlates of lexical ambiguity suppression mechanisms.     

Prioritizing new over old: an fMRI study of the preview search task

In visual search, observers can successfully ignore temporally separated distractors that are presented as a preview before onset of the search display. Previous behavioral studies have demonstrated the involvement of top-down selection mechanisms in preview search, biasing attention against the old set in favor of the more relevant new set. Using functional magnetic resonance imaging, we replicate and extend findings showing the involvement of superior and inferior parietal areas in the preview task when compared to both a relatively easy single-set search task and a more effortful full-set search task. In contrast, the effortful full-set search showed activation in the dorsolateral prefrontal cortex when compared to the single-set search, suggesting that this area is involved in rejecting additional distractors that could not be separated in time.     

Recognition memory for emotionally negative and neutral words: an ERP study

Scalp recorded event-related potentials were used to investigate the neural activity elicited by emotionally negative and emotionally neutral words during the performance of a recognition memory task. Behaviourally, the principal difference between the two word classes was that the false alarm rate for negative items was approximately double that for the neutral words. Correct recognition of neutral words was associated with three topographically distinct ERP memory 'old/new' effects: an early, bilateral, frontal effect which is hypothesised to reflect familiarity-driven recognition memory; a subsequent left parietally distributed effect thought to reflect recollection of the prior study episode; and a late onsetting, right-frontally distributed effect held to be a reflection of post-retrieval monitoring. The old/new effects elicited by negative words were qualitatively indistinguishable from those elicited by neutral items and, in the case of the early frontal effect, of equivalent magnitude also. However, the left parietal effect for negative words was smaller in magnitude and shorter in duration than that elicited by neutral words, whereas the right frontal effect was not evident in the ERPs to negative items. These differences between neutral and negative words in the magnitude of the left parietal and right frontal effects were largely attributable to the increased positivity of the ERPs elicited by new negative items relative to the new neutral items. Together, the behavioural and ERP findings add weight to the view that emotionally valenced words influence recognition memory primarily by virtue of their high levels of 'semantic cohesion', which leads to a tendency for 'false recollection' of unstudied items.     

Attention mechanisms in visual search -- an fMRI study

The human visual system is usually confronted with many different objects at a time, with only some of them reaching consciousness. Reaction-time studies have revealed two different strategies by which objects are selected for further processing: an automatic, efficient search process, and a conscious, so-called inefficient search [Treisman, A. (1991). Search, similarity, and integration of features between and within dimensions. Journal of Experimental Psychology: Human Perception and Performance, 17, 652--676; Treisman, A., & Gelade, G. (1980). A feature integration theory of attention. Cognitive Psychology, 12, 97--136; Wolfe, J. M. (1996). Visual search. In H. Pashler (Ed.), Attention. London: University College London Press]. Two different theories have been proposed to account for these search processes. Parallel theories presume that both types of search are treated by a single mechanism that is modulated by attentional and computational demands. Serial theories, in contrast, propose that parallel processing may underlie efficient search, but inefficient searching requires an additional serial mechanism, an attentional "spotlight" (Treisman, A., 1991) that successively shifts attention to different locations in the visual field. Using functional magnetic resonance imaging (fMRI), we show that the cerebral networks involved in efficient and inefficient search overlap almost completely. Only the superior frontal region, known to be involved in working memory [Courtney, S. M., Petit, L., Maisog, J. M., Ungerleider, L. G., & Haxby, J. V. (1998). An area specialized for spatial working memory in human frontal cortex. Science, 279, 1347--1351], and distinct from the frontal eye fields, that control spatial shifts of attention, was specifically involved in inefficient search. Activity modulations correlated with subjects' behavior best in the extrastriate cortical areas, where the amount of activity depended on the number of distracting elements in the display. Such a correlation was not observed in the parietal and frontal regions, usually assumed as being involved in spatial attention processing. These results can be interpreted in two ways: the most likely is that visual search does not require serial processing, otherwise we must assume the existence of a serial searchlight that operates in the extrastriate cortex but differs from the visuospatial shifts of attention involving the parietal and frontal regions.     

Recognition of emotional prosody and verbal components of spoken language: an fMRI study

This study examined the neural areas involved in the recognition of both emotional prosody and phonemic components of words expressed in spoken language using echo-planar, functional magnetic resonance imaging (fMRI). Ten right-handed males were asked to discriminate words based on either expressed emotional tone (angry, happy, sad, or neutral) or phonemic characteristics, specifically, initial consonant sound (bower, dower, power, or tower). Significant bilateral activity was observed in the detection of both emotional and verbal aspects of language when compared to baseline activity. We found that the detection of emotion compared with verbal detection resulted in significant activity in the right inferior frontal lobe. Conversely, the detection of verbal stimuli compared with the detection of emotion activated left inferior frontal lobe regions most significantly. Specific analysis of the anterior auditory cortex revealed increased right hemisphere activity during the detection of emotion compared to activity during verbal detection. These findings illustrate bilateral involvement in the detection of emotion in language while concomitantly showing significantly lateralized activity in both emotional and verbal detection, in both the temporal and frontal lobes.     

Impact of voice on emotional judgment of faces: an event-related fMRI study

Emotional information can be conveyed by various means of communication, such as propositional content, speech intonation, facial expression, and gestures. Prior studies have demonstrated that inputs from one modality can alter perception in another modality. To evaluate the impact of emotional intonation on ratings of emotional faces, a behavioral study first was carried out. Second, functional magnetic resonance (fMRI) was used to identify brain regions that mediate crossmodal effects of emotional prosody on judgments of facial expressions. In the behavioral study, subjects rated fearful and neutral facial expressions as being more fearful when accompanied by a fearful voice as compared to the same facial expressions without concomitant auditory stimulus, whereas no such influence on rating of faces was found for happy voices. In the fMRI experiment, this shift in rating of facial expressions in presence of a fearfully spoken sentence was correlated with the hemodynamic response in the left amygdala extending into the periamygdaloid cortex, which suggests that crossmodal effects on cognitive judgments of emotional information are mediated via these neuronal structures. Furthermore, significantly stronger activations were found in the mid-portion of the right fusiform gyrus during judgment of facial expressions in presence of fearful as compared to happy intonations, indicating that enhanced processing of faces within this region can be induced by the presence of threat-related information perceived via the auditory modality. Presumably, these increased extrastriate activations correspond to enhanced alertness, whereas responses within the left amygdala modulate cognitive evaluation of emotional facial expressions.     

Effects of aging on recognition of intentionally and incidentally stored words: an fMRI study

Functional magnetic resonance imaging (fMRI) was used to gain more insight in the mechanism underlying a decline in recognition memory function with age. Twelve young (23-27 years) and 12 older (63-67 years) healthy men performed two types of word encoding tasks, in which words were either incidentally or intentionally encoded for storage in memory. After a 30min delay, participants performed a recognition task. Older participants were less accurate and slower than young on the recognition task. In the both groups, successful retrieval was accompanied by activation in the left inferior frontal gyrus, left precentral gyrus and right cerebellum. Older participants showed additional activity in the bilateral medial prefrontal gyrus and right parahippocampal gyrus. Correlational analyses showed that only the additional parahippocampal activation correlated positively with task performance in the older but not young participants, suggesting that activation in this area served the purpose of functional compensation. The additional activation in the medial prefrontal cortex, on the other hand, was explained in terms of increased conflict, that is, reduced distinction between target and distracter words leading to increased simultaneous activity of both response tendencies. In a comparison between incidentally and intentionally remembered words the young group showed additional activation in the right middle occipital gyrus. This last result was explained in terms of strategic differences between the young and older group.