The experience of embarrassment provides a highly salient cue for the human moral apparatus. Interestingly, people also experience embarrassment on behalf of others' inappropriate conditions. The perceiver's embarrassment often lacks an equivalent expression of embarrassment in the social counterpart. The present study examines this phenomenon and distinguishes neural circuits involved in embarrassment with and embarrassment for another person's mishaps. Using functional magnetic resonance imaging, we show that the embarrassment on behalf of others engages the temporal pole and the medial prefrontal cortex, central structures of the mentalizing network, together with the anterior insula and anterior cingulate cortex. In contrast, sharing others' embarrassment additionally stimulated the posterior superior temporal sulcus (STS), which exhibited increased functional integration with inferior parietal and insular cortex areas. These findings characterize common neural circuits involved in the embodied representation of embarrassment and further unravel the unique role of the posterior STS in sharing others' affective state.
The aim of this Hypothesis and Theory is to question the recently increasing use of the "race" concept in contemporary genetic, psychiatric, neuroscience as well as social studies. We discuss "race" and related terms used to assign individuals to distinct groups and caution that also concepts such as "ethnicity" or "culture" unduly neglect diversity. We suggest that one factor contributing to the dangerous nature of the "race" concept is that it is based on a mixture of traditional stereotypes about "physiognomy", which are deeply imbued by colonial traditions. Furthermore, the social impact of "race classifications" will be critically reflected. We then examine current ways to apply the term "culture" and caution that while originally derived from a fundamentally different background, "culture" is all too often used as a proxy for "race", particularly when referring to the population of a certain national state or wider region. When used in such contexts, suggesting that all inhabitants of a geographical or political unit belong to a certain "culture" tends to ignore diversity and to suggest a homogeneity, which consciously or unconsciously appears to extend into the realm of biological similarities and differences. Finally, we discuss alternative approaches and their respective relevance to biological and cultural studies.
Rapid eye movement (REM) dreaming results in "emotionally intelligent encoding," according to the target article. Building on this, we argue that elaborative encoding alters emotional processing of upcoming events and thereby functions as prospective emotion regulation. After elaborative encoding, future events are appraised differently and result in a redirected emotional response. Disturbed elaborative encoding might be relevant for emotional dysregulation in psychopathology.
Second-person neuroscience offers a framework for the study of social emotions, such as embarrassment and pride. However, we propose that an enduring mental representation of oneself in relation to others without a continuous direct social interaction is possible. We call this state "social immersion" and will explain its impact on the neuroscience of social emotions.
The penetrance of genetic variation has been assumed to be higher at the level of neural phenotypes than at the level of behavioral phenotypes. One of the few attempts to validate this assumption is the study of Rose and Donohoe published in this issue. In this article, we will address 2 methodological issues we believe have to be considered for a better understanding of the present results. We briefly discuss potential solutions that might also help improve future meta-analyses of effect sizes in neuroimaging data.
The affective responses to another persons condition depend on the ability to reflect about anothers thoughts and intentions. This is relevant also for high-functioning individuals with ASD who have considerable difficulties in reading the intentions of others. With the present study we introduce a novel paradigm to induce vicarious embarrassment as a form of social pain. We predicted that the vicarious embarrassment experiences of high-functioning individuals with ASD should specifically decline in social contexts that require reflecting on anothers intentions. Thirty-two young adults with high-functioning ASD were matched with regards to age, gender, and verbal IQ to a control group. Vicarious embarrassment was examined with previously validated stimuli describing 30 situations that elicit vicarious embarrassment in the observer. The situations manipulated whether the displayed protagonist either accidentally or intentionally transgressed a social norm in public and participants rated their vicarious embarrassment from the observers perspective. The ASD group showed comparable vicarious embarrassment experience in response to observing anothers accidental norm transgressions but significantly reduced vicarious embarrassment when observing another who intentionally violated socials norms. Vicarious embarrassment was significantly correlated with trait empathy in the ASD group. In complex social scenarios individuals with ASD are impaired in reporting experience of vicarious embarrassment, primarily when it is required to reflect on anothers intentions. The present study thus contributes to a better understanding of how persons with ASD are affected in the diversity of empathic processes in the social, everyday life environment they are embedded in.
The mechanisms underlying hemispheric specialization of memory are not completely understood. Functional magnetic resonance imaging (fMRI) can be used to develop and test models of hemispheric specialization. In particular for memory tasks however, the interpretation of fMRI results is often hampered by the low reliability of the data. In the present study we therefore analyzed the test-retest reliability of fMRI brain activation related to an implicit memory encoding task, with a particular focus on brain activity of the medial temporal lobe (MTL). Fifteen healthy subjects were scanned with fMRI on two sessions (average retest interval 35?days) using a commonly applied novelty encoding paradigm contrasting known and unknown stimuli. To assess brain lateralization, we used three different stimuli classes that differed in their verbalizability (words, scenes, fractals). Test-retest reliability of fMRI brain activation was assessed by an intraclass-correlation coefficient (ICC), describing the stability of inter-individual differences in the brain activation magnitude over time. We found as expected a left-lateralized brain activation network for the words paradigm, a bilateral network for the scenes paradigm, and predominantly right-hemispheric brain activation for the fractals paradigm. Although these networks were consistently activated in both sessions on the group level, across-subject reliabilities were only poor to fair (ICCs???0.45). Overall, the highest ICC values were obtained for the scenes paradigm, but only in strongly activated brain regions. In particular the reliability of brain activity of the MTL was poor for all paradigms. In conclusion, for novelty encoding paradigms the interpretation of fMRI results on a single subject level is hampered by its low reliability. More studies are needed to optimize the retest reliability of fMRI activation for memory tasks.
Theory of Mind (ToM) is the ability to infer other peoples mental states like intentions or desires. ToM can be differentiated into affective (i.e., recognizing the feelings of another person) and cognitive (i.e., inferring the mental state of the counterpart) subcomponents. Recently, subcortical structures such as the basal ganglia (BG) have also been ascribed to the multifaceted concept ToM and most BG disorders have been reported to elicit ToM deficits. In order to assess both the correlates of affective and cognitive ToM as well as involvement of the basal ganglia, 30 healthy participants underwent event-related fMRI scanning, neuropsychological testing, and filled in questionnaires concerning different aspects of ToM and empathy. Directly contrasting affective (aff) as well as cognitive (cog) ToM to the control (phy) condition, activation was found in classical ToM regions, namely parts of the temporal lobe including the superior temporal sulcus, the supplementary motor area, and parietal structures in the right hemisphere. The contrast aff > phy yielded additional activation in the orbitofrontal cortex on the right and the cingulate cortex, the precentral and inferior frontal gyrus and the cerebellum on the left. The right BG were recruited in this contrast as well. The direct contrast aff > cog showed activation in the temporoparietal junction and the cingulate cortex on the right as well as in the left supplementary motor area. The reverse contrast cog > aff however did not yield any significant clusters. In summary, affective and cognitive ToM partly share neural correlates but can also be differentiated anatomically. Furthermore, the BG are involved in affective ToM and thus their contribution is discussed as possibly providing a motor component of simulation processes, particularly in affective ToM.
In the introduction to the special issue "The Neural Underpinnings of Vicarious Experience" the editors state that one "may feel embarrassed when witnessing another making a social faux pas". In our commentary we address this statement and ask whether this example introduces a vicarious or an empathic form of embarrassment. We elaborate commonalities and differences between these two forms of emotional experiences and discuss their underlying mechanisms. We suggest that both, vicarious and empathic emotions, originate from the simulation processes mirroring and mentalizing that depend on anchoring and adjustment. We claim the term "empathic emotion" to be reserved exclusively for incidents where perceivers and social targets have shared affective experience, whereas "vicarious emotion" offers a wider scope and also includes non-shared affective experiences. Both are supposed to be highly functional in social interactions.
The N-methyl-D-aspartate receptor (NMDAR) has been implicated in the pathophysiology of schizophrenia. Administered to healthy individuals, a subanesthetic dose of the noncompetitive NMDAR antagonist ketamine reproduces several psychopathological symptoms commonly observed in patients with schizophrenia. In a counterbalanced, placebo-controlled, double-blind, within-participants study, fifteen healthy subjects were administered a continuous subanesthetic S-ketamine infusion while cortical activation was measured using functional magnetic resonance imaging. While being scanned, subjects performed an overt word generation task. Ketamine-induced psychopathological symptoms were assessed with the Positive and Negative Syndrome Scale (PANSS). Ketamine administration elicited effects on psychopathology, including difficulties in abstract thinking, lack of spontaneity and flow of conversation as well as formal thought disorder. On a behavioral level, verbal fluency performance was unaffected. The PANSS score for formal thought disorder positively correlated with activation measures encompassing the left superior temporal gyrus, the right middle and inferior frontal gyrus and the precuneus. Difficulty in abstract thinking was correlated with pronounced activations in prefrontal as well as in anterior cingulate regions, whereas hyperactivations in the left superior temporal gyrus were found in association with a lack of spontaneity and flow of conversation. In the absence of behavioral impairments during verbal fluency, NMDAR blocking evoked psychopathological symptoms and cortical activations in regions previously reported in schizophrenia patients. The results provide further support for the hypothesis of an NMDAR dysfunction in the pathophysiology of schizophrenia.
Neurogranin (NRGN) is the main postsynaptic protein regulating the availability of calmodulin-Ca(2+) in neurons. NRGN is expressed exclusively in the brain, particularly in dendritic spines and has been implicated in spatial learning and hippocampal plasticity. Genetic variation in rs12807809 in the NRGN gene has recently been confirmed to be associated with schizophrenia in a meta-analysis of genome-wide association studies: the T-allele was found to be genome-wide significantly associated with schizophrenia. Cognitive tests and personality questionnaires were administered in a large sample of healthy subjects. Brain activation was measured with functional magnetic resonance imaging (fMRI) during an episodic memory encoding and retrieval task in a subsample. All subjects were genotyped for NRGN rs12807809. There was no effect of genotype on personality or cognitive measures in the large sample. Homozygote carriers of the T-allele showed better performance in the retrieval task during fMRI. After controlling for memory performance, differential brain activation was evident in the anterior cingulate cortex for the encoding and posterior cingulate regions during retrieval. We could demonstrate that rs12807809 of NRGN is associated with differential neural functioning in the anterior and posterior cingulate. These areas are involved in episodic memory processes and have been implicated in the pathophysiology of schizophrenia in structural and functional imaging as well as postmortem studies.
Functional magnetic resonance imaging (fMRI) can be combined with genotype assessment to identify brain systems that mediate genetic vulnerability to mental disorders ("imaging genetics"). A data analysis approach that is widely applied is "functional connectivity". In this approach, the temporal correlation between the fMRI signal from a pre-defined brain region (the so-called "seed point") and other brain voxels is determined. In this technical note, we show how the choice of freely selectable data analysis parameters strongly influences the assessment of the genetic modulation of connectivity features. In our data analysis we exemplarily focus on three methodological parameters: (i) seed voxel selection, (ii) noise reduction algorithms, and (iii) use of additional second level covariates. Our results show that even small variations in the implementation of a functional connectivity analysis can have an impact on the connectivity pattern that is as strong as the potential modulation by genetic allele variants. Some effects of genetic variation can only be found for one specific implementation of the connectivity analysis. A reoccurring difficulty in the field of psychiatric genetics is the non-replication of initially promising findings, partly caused by the small effects of single genes. The replication of imaging genetic results is therefore crucial for the long-term assessment of genetic effects on neural connectivity parameters. For a meaningful comparison of imaging genetics studies however, it is therefore necessary to provide more details on specific methodological parameters (e.g., seed voxel distribution) and to give information how robust effects are across the choice of methodological parameters.
Genome-wide association studies identified the single nucleotide polymorphism rs1344706 in ZNF804A as a common risk-variant for schizophrenia and bipolar disorder. Whereas the molecular function of ZNF804A is yet unclear, recent imaging genetics studies have started to characterize the neural systems architecture linking rs1344706 genotype to psychosis. Carring rs1344706 risk-alleles was associated with a decrease in functional connectivity within the dorsolateral prefrontal cortices (DLPFCs) as well as an increase in connectivity between the DLPFC and the hippocampal formation (HF) in the context of a working memory task. The present study aimed at replicating these findings in an independent sample of 94 healthy subjects. Subjects were genotyped for rs1344706 and performed a working memory task during functional magnetic resonance imaging. Results indicate no support for a decrease of functional coupling between the bilateral DLPFCs at higher ZNF804A risk status. However, the current data show the previously described alteration in functional coupling between the right DLPFC and the HFs, albeit with weaker effects. Decoupled by default, the functional connectivity between the right DLPFC and anterior HFs increased with the number of rs1344706 risk alleles. The present data support fronto-hippocampal dysconnectivity as intermediate phenotype linking rs1344706 genotype to psychosis. We discuss the issues in replicating the interhemispheric DLPFC coupling in light of the effect sizes rs1344706 genotype has on brain function, concluding that further independent replication studies are fundamentally needed to ascertain the role of rs1344706 in the functional integration of neural systems.
Ten years ago, neuroscientists began to study cultural phenomena by using functional MRI. Since then the number of publications in this field, termed cultural neuroscience (CN), has tremendously increased. In these studies, particular concepts of culture are implied, but rarely explicitly discussed. We argue that it is necessary to make these concepts a topic of debate in order to unravel the foundations of CN. From 40 fMRI studies we extracted two strands of reasoning: models investigating universal mechanisms for the formation of cultural groups and habits and, models assessing differences in characteristics among cultural groups. Both strands simplify culture as an inflexible set of traits and specificities. We question this rigid understanding of culture and highlight its hidden evaluative nature.
Rhyming words, as in songs or poems, is a universal feature of human language across all ages. In the present fMRI study a novel overt rhyming task was applied to determine the neural correlates of rhyme production. Fifteen right-handed healthy male volunteers participated in this verbal fluency study. Participants were instructed to overtly articulate as many words as possible either to a given initial letter (LVF) or to a semantic category (SVF). During the rhyming verbal fluency task (RVF), participants had to generate words that rhymed with pseudoword stimuli. On-line overt verbal responses were audiotaped in order to correct the imaging results for the number of generated words. Fewer words were generated in the rhyming compared to both the lexical and the semantic condition. On a neural level, all language tasks activated a language network encompassing the left inferior frontal gyrus, the middle and superior temporal gyri as well as the contralateral right cerebellum. Rhyming verbal fluency compared to both lexical and semantic verbal fluency demonstrated significantly stronger activation of left inferior parietal region. Generating novel rhyme words seems to be mainly mediated by the left inferior parietal lobe, a region previously found to be associated with meta-phonological as well as sub-lexical linguistic processes.
People vicariously experience embarrassment when observing others public pratfalls or etiquette violations. In two consecutive studies we investigated the subjective experience and the neural correlates of vicarious embarrassment for others in a broad range of situations. We demonstrated, first, that vicarious embarrassment was experienced regardless of whether the observed protagonist acted accidentally or intentionally and was aware or unaware that he/she was in an embarrassing situation. Second, using functional magnetic resonance imaging (fMRI), we showed that the anterior cingulate cortex and the left anterior insula, two cortical structures typically involved in vicarious feelings of others pain, are also strongly implicated in experiencing the social pain for others flaws and pratfalls. This holds true even for situations that engage protagonists not aware of their current predicament. Importantly, the activity in the anterior cingulate cortex and the left anterior insula positively correlated with individual differences in trait empathy. The present findings establish the empathic process as a fundamental prerequisite for vicarious embarrassment experiences, thus connecting affect and cognition to interpersonal processes."When we are living with people who have a delicate sense of propriety, we are in misery on their account when anything unbecoming is committed. So I always feel for and with Charlotte when a person is tipping his chair. She cannot endure it." [Elective Affinities, J. W. Goethe].
The production of language is one of the most complex and amazing skills in humans. Increasing evidence demonstrated that associative relations (e.g., car - garage) play an important role during concept formation but during speech production the effects and processing of associations are highly debated. Hence, the present study investigated the impact of associations and different SOAs on the production of sentences (Experiment 1) and on naming objects (Experiment 2). In an adapted version of the picture-word interference task, participants were asked to name two pictures using a standardized sentence (e.g., "The car is to the left of the trousers"). Thereby, a simultaneous (SOA is 0 ms) or slightly preceding (SOA is -150 ms) auditory or visual distractor had to be ignored. Distractors were related to the first noun (for example: "The car is to the left to the trousers", distractor: "garage") or to the second noun (distractor: "belt") or unrelated to both nouns (distractor: "bottle") of the sentence. At simultaneous presentation, visual and auditory distractors related to the first noun of the sentence prolonged naming responses (i.e., interference). For slightly preceding distractors, only visual presentation induced interference for the first noun of the sentence. During no condition, longer naming responses were found for the second noun of the sentence. These effects suggest that associatively related concepts are active during speech production and can be competitors, i.e., they lead to semantic interference. In Experiment 2, subjects had to name an object (e.g., car) while ignoring a visually presented distractor (e.g., motor). The stimulus set was the same as in Experiment 1. The results showed a facilitation effect if the distractor and the target were associatively related. Overall, the current results provide new insight in the models of speech production: while during single word production, associations facilitate naming, they interfere during sentence production. Hence, associations have an important influence on producing speech but the impact is varied by the context, i.e., single word or sentential.
The glutamatergic N-methyl-D-aspartate (NMDA) receptor has been implicated in the pathophysiology of schizophrenia. Administered to healthy volunteers, a subanesthetic dose of the non-competitive NMDA receptor antagonist ketamine leads to psychopathological symptoms similar to those observed in schizophrenia. In patients with schizophrenia, ketamine exacerbates the core symptoms of illness, supporting the hypothesis of a glutamatergic dysfunction. In a counterbalanced, placebo-controlled, double-blind study design, healthy subjects were administered a continuous subanesthetic S-ketamine infusion while differences in BOLD responses measured with fMRI were detected. During the scanning period, subjects performed continuous overt verbal fluency tasks (phonological, lexical and semantic). Ketamine-induced psychopathological symptoms were assessed with the Positive and Negative Syndrome Scale (PANSS). Ketamine elicited psychosis like psychopathology. Post-hoc t-tests revealed significant differences between placebo and ketamine for the amounts of words generated during lexical and semantic verbal fluency, while the phonological domain remained unaffected. Ketamine led to enhanced cortical activations in supramarginal and frontal brain regions for phonological and lexical verbal fluency, but not for semantic verbal fluency. Ketamine induces activation changes in healthy subjects similar to those observed in patients with schizophrenia, particularly in frontal and temporal brain regions. Our results provide further support for the hypothesis of an NMDA receptor dysfunction in the pathophysiology of schizophrenia.
The objective of this short review is to highlight rewarding aspects of social interactions for humans and discuss their neural basis. Thereby we report recent research findings to illustrate how social stimuli in general are processed in the reward system and highlight the role of Theory of Mind as one mediating process for experiencing social reward during social interactions. In conclusion we discuss clinical implications for psychiatry and psychotherapy.
Attention deficits belong to the main cognitive symptoms of schizophrenia and come along with altered neural activity in previously described cerebral networks. Given the high heritability of schizophrenia the question arises if impaired function of these networks is modulated by susceptibility genes and detectable in healthy risk allele carriers.
Memory dysfunction is a prominent feature in schizophrenia. Impairments of declarative memory have been consistently linked to alterations especially within hippocampal-prefrontal regions. Due to the high heritability of schizophrenia, susceptibility genes and their modulatory impact on the neural correlates on memory are of major relevance. In the present study the influence of the COMT val(158)met status on the neural correlates of declarative memory was investigated in healthy subjects.
Polymorphisms in the G72 (also named d-amino acid oxidase activator, DAOA) gene increase the vulnerability for schizophrenia and affective psychosis. Three recent genetic neuroimaging studies showed that variation in G72 influences the brain activity in the medial temporal lobe (MTL), supporting the hypothesis that G72 might play a modulatory role on brain activity in MTL structures. In the present study we therefore investigated the effect of G72 on the neural correlates of long-term memory encoding and retrieval in a large sample of healthy subjects (n=83) using functional magnetic resonance imaging. A face encoding and a face retrieval memory task were chosen because on the one hand they specifically activate MTL structures and on the other hand they tap into memory processes that are compromised in patients with schizophrenia and affective disorder. Despite a strong a-priori hypothesis of genotype group activation differences in the MTL along with a large sample size we did neither find an effect of G72 genotype status on brain activity in the MTL nor in any other brain regions. The present data therefore do not support the view of a general modulatory role of G72 on MTL brain activity, at least not in the domain of long-term memory encoding and retrieval. Our results highlight the importance of replication studies in genetic neuroimaging.
Human behaviour is generally guided by the anticipation of potential outcomes that are considered to be rewarding. Reward processing can thus be dissected into a phase of reward anticipation and a phase of reward consumption. A number of brain structures have been suggested to be involved in reward processing. However, it is unclear whether anticipation and consumption are mediated by the same or different neural networks. We examined the neural basis of these processes using functional magnetic resonance imaging (fMRI) in an incentive delay task offering either money or social approval. In both conditions participants (N=28) were given a cue indicating potential reward. In order to receive reward a target button had to be pushed within a certain time window (adapted for individual reaction time). Cues triggering either monetary or social reward anticipation were presented sessionwise. Imaging was performed on a 1.5-Tesla Philips scanner in an event-related design. Anticipation of both reward types activated brain structures constituting the brain reward system including the ventral striatum. In contrast to the task independent activity in the anticipation phase, reward consumption evoked different patterns of activation for money and social approval, respectively. While social stimuli were mainly associated with amygdala activation, the thalamus was more strongly activated by the presentation of monetary rewards. Our results identify dissociable neural networks for the anticipation and consumption of reward. The findings implicate that the neural mechanisms underlying reward consumption are more modality-specific than those for reward anticipation, and that they are mediated by subjective reward value.
Neuregulin 1 (NRG1) has been found to be associated with schizophrenia. Impaired performance in episodic memory tasks is an often replicated finding in this disorder. In functional neuroimaging studies, this dysfunction has been linked to signal changes in prefrontal and medial temporal areas. Therefore, it is of interest whether genes associated with the disorder, such as NRG1, modulate episodic memory performance and its neural correlates. Ninety-four healthy individuals performed an episodic memory encoding and a retrieval task while brain activation was measured with functional MRI. All subjects were genotyped for the single nucleotide polymorphism (SNP) rs35753505 in the NRG1 gene. The effect of genotype on brain activation was assessed with fMRI during the two tasks. While there were no differences in performance, brain activation in the cingulate gyrus (BA 24), the left middle frontal gyrus (BA 9), the bilateral fusiform gyrus and the left middle occipital gyrus (BA 19) was positively correlated with the number of risk alleles in NRG1 during encoding. During retrieval brain activation was positively correlated with the number of risk alleles in the left middle occipital gyrus (BA 19). NRG1 genotype does modulate brain activation during episodic memory processing in key areas for memory encoding and retrieval. The results suggest that subjects with risk alleles show hyperactivations in areas associated with elaborate encoding strategies.
Recent genetic studies found the A allele of the variant rs1006737 in the alpha 1C subunit of the L-type voltage-gated calcium channel (CACNA1C) gene to be overrepresented in patients suffering from bipolar disorder, schizophrenia or major depression. While the functions underlying the pathophysiology of these psychiatric disorders are yet unknown, impaired performance in verbal fluency tasks is an often replicated finding. We investigated the influence of the rs1006737 single nucleotide polymorphism (SNP) on verbal fluency and its neural correlates.
In our everyday life we process information from different modalities simultaneously with great ease. With the current study we had the following goals: to detect the neural correlates of (1) automatic semantic processing of associates and (2) to investigate the influence of different visual modalities on semantic processing. Stimuli were presented with a short SOA (350 ms) as subjects performed a lexical decision task. To minimize the variance and increase homogeneity within our sample, only male subjects were measured. Three experimental conditions were compared while brain activation was measured with a 3 T fMRI scanner: related word-pairs (e.g., frame-picture), unrelated word-pairs (e.g., frame-car) as well as word-nonword pairs (e.g., frame-fubber). They were presented uni- (word-word) and cross-modally (picture-word). Behavioral data revealed a priming effect for cross-modal and unimodal word-pairs. On a neural level, the unimodal condition revealed response suppression in bilateral fronto-parietal regions. Cross-modal priming led to response suppression within the right inferior frontal gyrus. Common areas of deactivation for both modalities were found in bilateral fronto-tempo-parietal regions. These results suggest that the processing of semantic associations presented in different modalities lead to modality-specific activation caused by early access routes. However, common activation for both modalities refers to a common neural network for semantic processing suggesting amodal processing.
"Lions" do not have "stripes". However, via the word "tiger" both words are closely connected within the semantic network. With the present study we pursued two goals: to detect neural correlates of (1) directly and indirectly related word pairs by means of priming, and (2) to assess the effect of presentation modality. Stimuli were presented with a short SOA of 350 ms as subjects performed a lexical decision task during fMRI measurement. Four experimental conditions were compared: directly related (picture-frame), indirectly related (anvil-nail), unrelated (steamboat-needle) and nonword trials (chalk-edan), presented in a uni- (word-word) and cross-modal (auditory-word) version. Behavioral data revealed a modality-independent priming effect only for direct semantic priming. On a neural level, directly linked words led to left-lateralized activations in fronto-temporo-parietal areas. Indirect priming led to right-hemispheric fronto-parietal signal changes. Common areas of activation for uni- and cross-modal priming were found within the left middle temporal gyrus and right precuneus for direct priming and within the right insula for indirect priming. The comparison of the semantic distances (direct>indirect) showed one region activated modality-independent: the precuneus. Direct priming is associated with activation clusters corresponding to a large left-lateralized network. Indirect priming recruits right-hemispheric regions, reflecting widespread semantic fields and attentional components. The modality-independent comparison of direct and indirect priming revealed common areas of activation supporting an amodal rather than multiple semantic systems. The activation related to semantic distances underpins the special role of the precuneus. This region is involved in semantic priming and association processing whereas episodic memory contents might be addressed.
Impaired performance in verbal fluency tasks is an often replicated finding in schizophrenia. In functional neuroimaging studies, this dysfunction has been linked to signal changes in prefrontal and temporal areas. Since schizophrenia has a high heritability, it is of interest whether susceptibility genes for the disorder, such as NRG1, modulate verbal fluency performance and its neural correlates. Four hundred twenty-nine healthy individuals performed a semantic and a lexical verbal fluency task. A subsample of 85 subjects performed an overt semantic verbal fluency task while brain activation was measured with functional magnetic resonance imaging (MRI). NRG1 (SNP8NRG221533; rs35753505) status was determined and correlated with verbal fluency performance and brain activation. For the behavioral measure, there was a linear effect of NRG1 status on semantic but not on lexical verbal fluency. Performance decreased with number of risk-alleles. In the fMRI experiment, decreased activation in the left inferior frontal and the right middle temporal gyri as well as the anterior cingulate gyrus was correlated with the number of risk-alleles in the semantic verbal fluency task. NRG1 genotype does influence language production on a semantic level in conjunction with the underlying neural systems. These findings are in line with results of studies in schizophrenia and may explain some of the cognitive and brain activation variation found in the disorder. More generally, NRG1 might be one of several genes that influence semantic language capacities.
Variation in the val(158)met polymorphism of the COMT gene has been found to be associated with cognitive performance. In functional neuroimaging studies, this dysfunction has been linked to signal changes in prefrontal areas. Given the complex modulation and functional heterogeneity of frontal lobe systems, further specification of COMT gene-related phenotypes differing in prefrontally mediated cognitive performance are of major interest. Eighty healthy individuals (54 men, 26 women; mean age 23.3 years) performed an overt semantic verbal fluency task while brain activation was measured with functional magnetic resonance imaging (fMRI). COMT val(158)met genotype was determined and correlated with brain activation measured with fMRI during the task. Although there were no differences in performance, brain activation in the left inferior frontal gyrus [Brodmann area 10] was positively correlated with the number of val alleles in the COMT gene. COMT val(158)met status modulates brain activation during the language production on a semantic level in an area related to executive functions.
The ability to segment continuous linguistic information online into larger, meaningful units is a key element in narrative comprehension. Narrative shifts, i.e. transitions between individual units, are postulated to continuously update the mental situation model. Their cerebral correlates, however, have hardly been investigated. Under highly naturalistic conditions this study seeks to identify the neural correlates of implicit processing of narrative shifts during continuous speech comprehension. 16 male native German speakers listened passively to a German novella for 23 min while BOLD signal was recorded with fMRI. Text comprehension was tested in a short post-scan interview asking for critical episodes of the story. Narrative shifts were defined on the basis of a macropropositional analysis. Compared to listening to text passages of the narrative that neither contained narrative shifts nor structurally similar linguistic control events (i.e., sentence boundaries), narrative shifts evoked increased BOLD signal changes in the right temporal gyrus, precuneus and posterior/middle cingulate cortex bilaterally. When narrative shifts were contrasted with sentence boundaries, activation in the right precuneus and cingulate cortex remained significant. The results strengthen the relevance of medial parietal structures for natural language comprehension. More precisely, the precuneus and posterior cingulate appear to be the neural substrate for updating mental story representations and can be regarded as critical parts of a more complex, distributed neural network underlying story comprehension.
In the last years, several susceptibility genes for psychiatric disorders have been identified, among others G72 (also named D-amino acid oxidase activator, DAOA). Typically, the high-risk variant of a vulnerability gene is associated with decreased cognitive functions already in healthy individuals. In a recent study however, a positive effect of the high-risk variant of G72 on verbal working memory was reported. In the present study, we therefore examined the relationship between G72 genotype status and a broad range of cognitive functions in 423 healthy individuals.
For successful interpersonal communication, inferring intentions, goals or desires of others is highly advantageous. Increasingly, humans also interact with computers or robots. In this study, we sought to determine to what degree an interactive task, which involves receiving feedback from social partners that can be used to infer intent, engaged the medial prefrontal cortex, a region previously associated with Theory of Mind processes among others. Participants were scanned using fMRI as they played an adapted version of the Prisoners Dilemma Game with alleged human and computer partners who were outside the scanner. The medial frontal cortex was activated when both human and computer partner were played, while the direct contrast revealed significantly stronger signal change during the human-human interaction. The results suggest a link between activity in the medial prefrontal cortex and the partner played in a mentalising task. This signal change was also present for to the computers partner. Implying agency or a will to non-human actors might be an innate human resource that could lead to an evolutionary advantage.
The ability to mentalize, i.e. develop a Theory of Mind (ToM), enables us to anticipate and build a model of the thoughts, emotions and intentions of others. It has long been hypothesised that women differ from men in their mentalizing abilities. In the present fMRI study we examined the impact of (1) gender (women vs. men) and (2) game partner (human vs. computer) on ToM associated neural activity in the medial prefrontal cortex. Groups of men (n = 12) and women (n = 12) interacted in an iterated classical prisoners dilemma forced choice situation with alleged human and computer partners who were outside the scanner.
Motivation for goal-directed behaviour largely depends on the expected value of the anticipated reward. The aim of the present study was to examine how different levels of reward value are coded in the brain for two common forms of human reward: money and social approval. To account for gender differences 16 male and 16 female participants performed an incentive delay task expecting to win either money or positive social feedback. fMRI recording during the anticipation phase revealed proportional activation of neural structures constituting the human reward system for increasing levels of reward, independent of incentive type. However, in men activation in the prospect of monetary rewards encompassed a wide network of mesolimbic brain regions compared to only limited activation for social rewards. In contrast, in women, anticipation of either incentive type activated identical brain regions. Our findings represent an important step towards a better understanding of motivated behaviour by taking into account individual differences in reward valuation.
G72 is a vulnerability gene for schizophrenia and affective psychosis, disorders that are characterized by deficits in working memory. In the present study we investigated whether the G72 genotype influences verbal and spatial working memory functions in healthy individuals. Working memory was assessed at the behavioural level in 423 subjects using the spatial span of the Wechsler Memory Scale (spatial working memory) and the letter-number-span test (verbal working memory). In a sub-sample of 83 subjects, we assessed working memory functions also at the neural level using functional magnetic resonance imaging during a classical letter variant of the n-back task. Unexpectedly the high risk allele carriers performed better in the verbal working memory task than the other subjects. These behavioural differences were accompanied by brain activation differences in the right parahippocampus, a brain region that plays a major role in schizophrenia and affective disorders. The high risk variant of a vulnerability gene therefore does not necessarily have to negatively affect cognitive abilities per se, but may even have beneficial effects on cognitive functions in the non-affected population.
Categorization is a basic principle of knowledge organization in the brain. The goal of the current fMRI study was to compare the neural correlates of thematic (e.g., car - garage) and taxonomic (e.g., couch - bed) categories under automatic processing conditions using auditory-to-visual semantic priming. Behavioral data revealed a priming effect for thematically but not for taxonomically related word pairs. On a neural level, thematically related words led to a left-lateralized temporal activation (superior temporal sulcus), whereas taxonomically related word pairs evoked a right-lateralized frontal activation and within the hippocampus. A direct comparison between both categories revealed enhanced activation for thematically related and response suppression for taxonomically related trials in the left superior temporal sulcus. These results suggest that processing of thematic and taxonomic categories leads to activation of distinct brain areas. The mainly right-lateralized fronto-temporal activation for taxonomic relations suggests increased attention and effort for processing this category. The interaction within the left superior temporal sulcus reflects the processing and retrieval of semantic relations whereby specific memory contents seem to influence the direction of activation.
We studied the somatovisceral response pattern of vicarious embarrassment for someone elses inappropriate condition. Participants (N=54) were confronted with hand-drawn sketches depicting public situations and were instructed to rate the intensity of their vicarious embarrassment. The inappropriate condition varied according to the attribution of intentionality (absent/present) and awareness (absent/present). Irrespective of these attributions, participants reported stronger vicarious embarrassment in comparison to neutral situations. Across a set of eleven somatovisceral variables vicarious embarrassment elicited a pattern of increased autonomic activation which was modulated by the awareness of the protagonist about the ongoing norm violation. The somatovisceral response pattern matches previous findings for the first-person experience of embarrassment. Together, these results support the hypothesis that processes of perspective taking also mediate the vicarious experience of embarrassment.
Crossed language dominance is a rare form of language lateralization, characterized by a dissociation of anterior and posterior language regions. We present the case of a healthy subject whose language lateralization pattern, as assessed by functional magnetic resonance imaging, is reliably characterized as crossed language dominance based on a word generation task, but typical left-lateralized when a semantic decision task is applied. A single language task is therefore not sufficient to characterize language lateralization, at least not for subjects with rare forms of language dominance. In the pre-surgical diagnostic of language lateralization, several language tasks tapping into different aspects of language functions should be applied.
Previous studies have shown overlapping neural activations for observation and execution or imitation of emotional facial expressions. These shared representations have been assumed to provide indirect evidence for a human mirror neuron system, which is suggested to be a prerequisite of action comprehension. We aimed at clarifying whether shared representations in and beyond human mirror areas are specifically activated by affective facial expressions or whether they are activated by facial expressions independent of the emotional meaning. During neuroimaging, participants observed and executed happy and non-emotional facial expressions. Shared representations were revealed for happy facial expressions in the pars opercularis, the precentral gyrus, in the superior temporal gyrus/medial temporal gyrus (MTG), in the pre-supplementary motor area and in the right amygdala. All areas showed less pronounced activation in the non-emotional condition. When directly compared, significant stronger neural responses emerged for happy facial expressions in the pre-supplementary motor area and in the MTG than for non-emotional stimuli. We assume that activation of shared representations depends on the affect and (social) relevance of the facial expression. The pre-supplementary motor area is a core-shared representation-structure supporting observation and execution of affective contagious facial expressions and might have a modulatory role during the preparation of executing happy facial expressions.
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