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1.  An Affective Circumplex Model of Neural Systems Subserving Valence, Arousal, & Cognitive Overlay During the Appraisal of Emotional Faces 
Neuropsychologia  2008;46(8):2129-2139.
Increasing evidence supports the existence of distinct neural systems that subserve two dimensions of affect – arousal and valence. Ten adult participants underwent functional magnetic resonance imaging during which they were presented a range of standardized faces and then asked, during the scan, to rate the emotional expressions of the faces along the dimensions of arousal and valence. Lower ratings of arousal accompanied greater activity in the amygdala complex, cerebellum, dorsal pons, and right medial prefrontal cortex. More negative ratings of valence accompanied greater activity in the dorsal anterior cingulate and parietal cortices. Extreme ratings of valence (highly positive and highly negative ratings) accompanied greater activity in the temporal cortex and fusiform gyrus. Building on an empirical literature which suggests that the amygdala serves as a salience and ambiguity detector, we interpret our findings as showing that a face rated has having low arousal is more ambiguous and a face rated as having extreme valence is more personally salient. This explains how both low arousal and extreme valence lead to greater activation of an ambiguity/salience system subserved by the amygdala, cerebellum, and dorsal pons. In addition, the right medial prefrontal cortex appears to down-regulate individual ratings of arousal, whereas the fusiform and related temporal cortices seem to up-regulate individual assessments of extreme valence when individual ratings are studied relative to group reference ratings for each stimulus. The simultaneous assessment of the effects of arousal and valence proved essential for the identification of neural systems contributing to the processing of emotional faces.
doi:10.1016/j.neuropsychologia.2008.02.032
PMCID: PMC2486369  PMID: 18440572
fMRI; Affect; Circumplex; Arousal; Valence; Face processing
2.  Lexical Tone Recognition with an Artificial Neural Network 
Ear and hearing  2008;29(3):326-335.
Objectives
Tone production is particularly important for communicating in tone languages such as Mandarin Chinese. In the present study, an artificial neural network was used to recognize tones produced by adult native speakers. The purposes of the study were (1) to test the sensitivity of the neural network to speaker variation typically in adult speaker groups, (2) to evaluate two normalization procedures to overcome the effects of speaker variation, and (3) to compare tone recognition performance of the neural network with that of the human listeners.
Design
A feedforward multilayer neural network was used. Twenty-nine adult native Mandarin Chinese speakers were recruited to record tone samples. The F0 contours of the vowel part of the 1044 monosyllabic words recorded were extracted using an autocorrelation method. Samples from the F0 contours were used as inputs to the neural network. The efficacy of the neural network was first tested by varying the number of inputs and the number of neurons in the hidden layer from 1 to 16. The sensitivity of the neural network to speaker variation was tested by (1) using the raw F0 data from speech tokens of a number of randomly drawn speakers that varied from 1 to 29, (2) using the raw F0 data from speech tokens of either male-only or female-only speakers, and (3) using two sets of normalized F0 data (i.e., tone 1-based normalization and first-order derivative) from speech tokens from a number of randomly drawn speakers that varied from 1 to 29. The recognition performance of the neural network under several experimental conditions was compared with the corresponding recognition performance of 10 normal-hearing, native Mandarin Chinese speaking adult listeners.
Results
Three inputs and four hidden neurons were found to be sufficient for the neural network to perform at about 85% correct using speech samples without normalization. The performance of the neural network was affected by variation across speakers particularly between genders. Using the tone 1-based normalization procedure, the performance of the neural network improved significantly. The recognition accuracy of the neural network as a whole or for each tone was comparable with that of the human listeners.
Conclusions
The neural network can be used to evaluate the tone production of Mandarin Chinese speaking adults with human listener-like recognition accuracy. The tone 1-based normalization procedure improves the performance of the neural network to human listener-like accuracy. The success of our neural network in recognizing tones from multiple speakers supports its utility for evaluating tone production. Further testing of the neural network with hearing-impaired speakers might reveal its potential use for clinical evaluation of tone production.
doi:10.1097/AUD.0b013e3181662c42
PMCID: PMC2562432  PMID: 18453884
3.  The Detection of Communicative Signals Directed at the Self in Infant Prefrontal Cortex 
A precondition for successful communication between people is the detection of signals indicating the intention to communicate, such as eye contact or calling a person's name. In adults, establishing communication by eye contact or calling a person's name results in overlapping activity in right prefrontal cortex, suggesting that, regardless of modality, the intention to communicate is detected by the same brain region. We measured prefrontal cortex responses in 5-month-olds using near-infrared spectroscopy (NIRS) to examine the neural basis of detecting communicative signals across modalities in early development. Infants watched human faces that either signaled eye contact or directed their gaze away from the infant, and they also listened to voices that addressed them with their own name or another name. The results revealed that infants recruit adjacent but non-overlapping regions in the left dorsal prefrontal cortex when they process eye contact and own name. Moreover, infants that responded sensitively to eye contact in the one prefrontal region were also more likely to respond sensitively to their own name in the adjacent prefrontal region as revealed in a correlation analysis, suggesting that responding to communicative signals in these two regions might be functionally related. These NIRS results suggest that infants selectively process and attend to communicative signals directed at them. However, unlike adults, infants do not seem to recruit a common prefrontal region when processing communicative signals of different modalities. The implications of these findings for our understanding of infants’ developing communicative abilities are discussed.
doi:10.3389/fnhum.2010.00201
PMCID: PMC2981376  PMID: 21088694
eye contact; name; communication; intention; prefrontal cortex; infancy
4.  Children’s and Adults’ Conceptualization and Evaluation of Lying and Truth-telling 
Infant and child development  2009;18(4):307-322.
The present study examined children’s and adults’ categorization and moral judgment of truthful and untruthful statements. 7-, 9-and 11-year-old Chinese children and college students read stories in which story characters made truthful or untruthful statements and were asked to classify and evaluate the statements. The statements varied in terms of whether the speaker intended to help or harm a listener and whether the statement was made in a setting that called for informational accuracy or politeness. Results showed that the communicative intent and setting factors jointly influence children’s categorization of lying and truth-telling, which extends an earlier finding (Lee & Ross, 1997) to childhood. Also, we found that children’s and adults’ moral judgments of lying and truth-telling were influenced by the communicative intent but not the setting factor. The present results were discussed in terms of Sweetser’s (1987) folkloristic model of lying.
doi:10.1002/icd.631
PMCID: PMC2951330  PMID: 20936049
lying; moral evaluation; white lies; deception; politeness; Chinese; culture
5.  Sensory-Cognitive Interaction in the Neural Encoding of Speech in Noise: A Review 
Background
Speech-in-noise (SIN) perception is one of the most complex tasks faced by listeners on a daily basis. Although listening in noise presents challenges for all listeners, background noise inordinately affects speech perception in older adults and in children with learning disabilities. Hearing thresholds are an important factor in SIN perception, but they are not the only factor. For successful comprehension, the listener must perceive and attend to relevant speech features, such as the pitch, timing, and timbre of the target speaker’s voice. Here, we review recent studies linking SIN and brainstem processing of speech sounds.
Purpose
To review recent work that has examined the ability of the auditory brainstem response to complex sounds (cABR), which reflects the nervous system’s transcription of pitch, timing, and timbre, to be used as an objective neural index for hearing-in-noise abilities.
Study Sample
We examined speech-evoked brainstem responses in a variety of populations, including children who are typically developing, children with language-based learning impairment, young adults, older adults, and auditory experts (i.e., musicians).
Data Collection and Analysis
In a number of studies, we recorded brainstem responses in quiet and babble noise conditions to the speech syllable /da/ in all age groups, as well as in a variable condition in children in which /da/ was presented in the context of seven other speech sounds. We also measured speech-in-noise perception using the Hearing-in-Noise Test (HINT) and the Quick Speech-in-Noise Test (QuickSIN).
Results
Children and adults with poor SIN perception have deficits in the subcortical spectrotemporal representation of speech, including low-frequency spectral magnitudes and the timing of transient response peaks. Furthermore, auditory expertise, as engendered by musical training, provides both behavioral and neural advantages for processing speech in noise.
Conclusions
These results have implications for future assessment and management strategies for young and old populations whose primary complaint is difficulty hearing in background noise. The cABR provides a clinically applicable metric for objective assessment of individuals with SIN deficits, for determination of the biologic nature of disorders affecting SIN perception, for evaluation of appropriate hearing aid algorithms, and for monitoring the efficacy of auditory remediation and training.
doi:10.3766/jaaa.21.9.3
PMCID: PMC3075209  PMID: 21241645
Auditory brainstem response; evoked potentials; frequency; musicians; speech in noise; timing
6.  Detecting Sarcasm from Paralinguistic Cues: Anatomic and Cognitive Correlates in Neurodegenerative Disease 
NeuroImage  2009;47(4):2005-2015.
While sarcasm can be conveyed solely through contextual cues such as counterfactual or echoic statements, face-to-face sarcastic speech may be characterized by specific paralinguistic features that alert the listener to interpret the utterance as ironic or critical, even in the absence of contextual information. We investigated the neuroanatomy underlying failure to understand sarcasm from dynamic vocal and facial paralinguistic cues. Ninety subjects (20 frontotemporal dementia, 11 semantic dementia [SemD], 4 progressive nonfluent aphasia, 27 Alzheimer’s disease, 6 corticobasal degeneration, 9 progressive supranuclear palsy, 13 healthy older controls) were tested using the Social Inference – Minimal subtest of The Awareness of Social Inference Test (TASIT). Subjects watched brief videos depicting sincere or sarcastic communication and answered yes-no questions about the speaker’s intended meaning. All groups interpreted Sincere (SIN) items normally, and only the SemD group was impaired on the Simple Sarcasm (SSR) condition. Patients failing the SSR performed more poorly on dynamic emotion recognition tasks and had more neuropsychiatric disturbances, but had better verbal and visuospatial working memory than patients who comprehended sarcasm. Voxel-based morphometry analysis of SSR scores in SPM5 demonstrated that poorer sarcasm comprehension was predicted by smaller volume in bilateral posterior parahippocampii (PHc), temporal poles, and R medial frontal pole (pFWE<0.05). This study provides lesion data suggesting that the PHc may be involved in recognizing a paralinguistic speech profile as abnormal, leading to interpretive processing by the temporal poles and right medial frontal pole that identifies the social context as sarcastic, and recognizes the speaker’s paradoxical intentions.
doi:10.1016/j.neuroimage.2009.05.077
PMCID: PMC2720152  PMID: 19501175
7.  Cortical Representations of Symbols, Objects, and Faces Are Pruned Back during Early Childhood 
Cerebral Cortex (New York, NY)  2010;21(1):191-199.
Regions of human ventral extrastriate visual cortex develop specializations for natural categories (e.g., faces) and cultural artifacts (e.g., words). In adults, category-based specializations manifest as greater neural responses in visual regions of the brain (e.g., fusiform gyrus) to some categories over others. However, few studies have examined how these specializations originate in the brains of children. Moreover, it is as yet unknown whether the development of visual specializations hinges on “increases” in the response to the preferred categories, “decreases” in the responses to nonpreferred categories, or “both.” This question is relevant to a long-standing debate concerning whether neural development is driven by building up or pruning back representations. To explore these questions, we measured patterns of visual activity in 4-year-old children for 4 categories (faces, letters, numbers, and shoes) using functional magnetic resonance imaging. We report 2 key findings regarding the development of visual categories in the brain: 1) the categories “faces” and “symbols” doubly dissociate in the fusiform gyrus before children can read and 2) the development of category-specific responses in young children depends on cortical responses to nonpreferred categories that decrease as preferred category knowledge is acquired.
doi:10.1093/cercor/bhq078
PMCID: PMC3000569  PMID: 20457691
development; fMRI; fusiform gyrus; pruning
8.  Stress and anxiety: Structural plasticity and epigenetic regulation as a consequence of stress 
Neuropharmacology  2011;62(1):3-12.
The brain is the central organ of stress and adaptation to stress because it perceives and determines what is threatening, as well as the behavioral and physiological responses to the stressor. The adult, as well as developing brain, possess a remarkable ability to show reversible structural and functional plasticity in response to stressful and other experiences, including neuronal replacement, dendritic remodeling, and synapse turnover. This is particularly evident in the hippocampus, where all three types of structural plasticity have been recognized and investigated, using a combination of morphological, molecular, pharmacological, electrophysiological and behavioral approaches. The amygdala and the prefrontal cortex, brain regions involved in anxiety and fear, mood, cognitive function and behavioral control, also show structural plasticity. Acute and chronic stress cause an imbalance of neural circuitry subserving cognition, decision making, anxiety and mood that can increase or decrease expression of those behaviors and behavioral states. In the short term, such as for increased fearful vigilance and anxiety in a threatening environment, these changes may be adaptive; but, if the danger passes and the behavioral state persists along with the changes in neural circuitry, such maladaptation may need intervention with a combination of pharmacological and behavioral therapies, as is the case for chronic or mood anxiety disorders. We shall review cellular and molecular mechanisms, as well as recent work on individual differences in anxiety-like behavior and also developmental influences that bias how the brain responds to stressors. Finally, we suggest that such an approach needs to be extended to other brain areas that are also involved in anxiety and mood.
doi:10.1016/j.neuropharm.2011.07.014
PMCID: PMC3196296  PMID: 21807003
9.  Children’s and adults’ neural bases of verbal and nonverbal ‘Theory of Mind’ 
Neuropsychologia  2007;45(7):1522-1532.
Theory of mind (ToM) - our ability to predict behaviors of others in terms of their underlying intentions - has been examined through verbal and nonverbal false-belief (FB) tasks. Previous brain imaging studies of ToM in adults have implicated medial prefrontal cortex (mPFC) and temporo-parietal junction (TPJ) for adults’ ToM ability. To examine age and modality related differences and similarities in neural correlates of ToM, we tested 16 adults (18-40 years-old) and 12 children (8-12 years-old) with verbal (story) and nonverbal (cartoon) FB tasks, using functional magnetic resonance imaging (fMRI). Both age groups showed significant activity in the TPJ bilaterally and right inferior parietal lobule (IPL) in a modality-independent manner, indicating that these areas are important for ToM during both adulthood and childhood, regardless of modality. We also found significant age-related differences in the ToM condition-specific activity for the story and cartoon tasks in the left inferior frontal gyrus (IFG) and left TPJ. These results suggest that depending on the modality adults may utilize different brain regions from children in understanding ToM.
doi:10.1016/j.neuropsychologia.2006.11.017
PMCID: PMC1868677  PMID: 17208260
fMRI; Theory of Mind; Cognitive Development; Language; Temporo-parietal junction
10.  The Neural Correlates of Emotional Prosody Comprehension: Disentangling Simple from Complex Emotion 
PLoS ONE  2011;6(12):e28701.
Background
Emotional prosody comprehension (EPC), the ability to interpret another person's feelings by listening to their tone of voice, is crucial for effective social communication. Previous studies assessing the neural correlates of EPC have found inconsistent results, particularly regarding the involvement of the medial prefrontal cortex (mPFC). It remained unclear whether the involvement of the mPFC is linked to an increased demand in socio-cognitive components of EPC such as mental state attribution and if basic perceptual processing of EPC can be performed without the contribution of this region.
Methods
fMRI was used to delineate neural activity during the perception of prosodic stimuli conveying simple and complex emotion. Emotional trials in general, as compared to neutral ones, activated a network comprising temporal and lateral frontal brain regions, while complex emotion trials specifically showed an additional involvement of the mPFC, premotor cortex, frontal operculum and left insula.
Conclusion
These results indicate that the mPFC and premotor areas might be associated, but are not crucial to EPC. However, the mPFC supports socio-cognitive skills necessary to interpret complex emotion such as inferring mental states. Additionally, the premotor cortex involvement may reflect the participation of the mirror neuron system for prosody processing particularly of complex emotion.
doi:10.1371/journal.pone.0028701
PMCID: PMC3236212  PMID: 22174872
11.  Can You Hear Me Now? Musical Training Shapes Functional Brain Networks for Selective Auditory Attention and Hearing Speech in Noise 
Even in the quietest of rooms, our senses are perpetually inundated by a barrage of sounds, requiring the auditory system to adapt to a variety of listening conditions in order to extract signals of interest (e.g., one speaker's voice amidst others). Brain networks that promote selective attention are thought to sharpen the neural encoding of a target signal, suppressing competing sounds and enhancing perceptual performance. Here, we ask: does musical training benefit cortical mechanisms that underlie selective attention to speech? To answer this question, we assessed the impact of selective auditory attention on cortical auditory-evoked response variability in musicians and non-musicians. Outcomes indicate strengthened brain networks for selective auditory attention in musicians in that musicians but not non-musicians demonstrate decreased prefrontal response variability with auditory attention. Results are interpreted in the context of previous work documenting perceptual and subcortical advantages in musicians for the hearing and neural encoding of speech in background noise. Musicians’ neural proficiency for selectively engaging and sustaining auditory attention to language indicates a potential benefit of music for auditory training. Given the importance of auditory attention for the development and maintenance of language-related skills, musical training may aid in the prevention, habilitation, and remediation of individuals with a wide range of attention-based language, listening and learning impairments.
doi:10.3389/fpsyg.2011.00113
PMCID: PMC3115514  PMID: 21716636
attention; cortical variability; auditory-evoked potentials; music; musicians; speech in noise; prefrontal cortex; language
12.  Abnormalities of Intrinsic Functional Connectivity in Autism Spectrum Disorders 
NeuroImage  2009;47(2):764-772.
Autism spectrum disorders (ASD) impact social functioning and communication, and individuals with these disorders often have restrictive and repetitive behaviors. Accumulating data indicate that ASD is associated with alterations of neural circuitry. Functional MRI (FMRI) studies have focused on connectivity in the context of psychological tasks. However, even in the absence of a task, the brain exhibits a high degree of functional connectivity, known as intrinsic or resting connectivity. Notably, the default network, which includes the posterior cingulate cortex, retro-splenial, lateral parietal cortex/angular gyrus, medial prefrontal cortex, superior frontal gyrus, temporal lobe, and parahippocampal gyrus, is strongly active when there is no task. Altered intrinsic connectivity within the default network may underlie offline processing that may actuate ASD impairments. Using FMRI, we sought to evaluate intrinsic connectivity within the default network in ASD. Relative to controls, the ASD group showed weaker connectivity between the posterior cingulate cortex and superior frontal gyrus and stronger connectivity between the posterior cingulate cortex and both the right temporal lobe and right parahippocampal gyrus. Moreover, poorer social functioning in the ASD group was correlated with weaker connectivity between the posterior cingulate cortex and the superior frontal gyrus. In addition, more severe restricted and repetitive behaviors in ASD were correlated with stronger connectivity between the posterior cingulate cortex and right parahippocampal gyrus. These findings indicate that ASD subjects show altered intrinsic connectivity within the default network, and connectivity between these structures is associated with specific ASD symptoms.
doi:10.1016/j.neuroimage.2009.04.069
PMCID: PMC2731579  PMID: 19409498
13.  Time Course of the Involvement of the Right Anterior Superior Temporal Gyrus and the Right Fronto-Parietal Operculum in Emotional Prosody Perception 
PLoS ONE  2008;3(5):e2244.
In verbal communication, not only the meaning of the words convey information, but also the tone of voice (prosody) conveys crucial information about the emotional state and intentions of others. In various studies right frontal and right temporal regions have been found to play a role in emotional prosody perception. Here, we used triple-pulse repetitive transcranial magnetic stimulation (rTMS) to shed light on the precise time course of involvement of the right anterior superior temporal gyrus and the right fronto-parietal operculum. We hypothesized that information would be processed in the right anterior superior temporal gyrus before being processed in the right fronto-parietal operculum. Right-handed healthy subjects performed an emotional prosody task. During listening to each sentence a triplet of TMS pulses was applied to one of the regions at one of six time points (400–1900 ms). Results showed a significant main effect of Time for right anterior superior temporal gyrus and right fronto-parietal operculum. The largest interference was observed half-way through the sentence. This effect was stronger for withdrawal emotions than for the approach emotion. A further experiment with the inclusion of an active control condition, TMS over the EEG site POz (midline parietal-occipital junction), revealed stronger effects at the fronto-parietal operculum and anterior superior temporal gyrus relative to the active control condition. No evidence was found for sequential processing of emotional prosodic information from right anterior superior temporal gyrus to the right fronto-parietal operculum, but the results revealed more parallel processing. Our results suggest that both right fronto-parietal operculum and right anterior superior temporal gyrus are critical for emotional prosody perception at a relatively late time period after sentence onset. This may reflect that emotional cues can still be ambiguous at the beginning of sentences, but become more apparent half-way through the sentence.
doi:10.1371/journal.pone.0002244
PMCID: PMC2373925  PMID: 18493307
14.  Neural foundations to moral reasoning and antisocial behavior 
A common feature of the antisocial, rule-breaking behavior that is central to criminal, violent and psychopathic individuals is the failure to follow moral guidelines. This review summarizes key findings from brain imaging research on both antisocial behavior and moral reasoning, and integrates these findings into a neural moral model of antisocial behavior. Key areas found to be functionally or structurally impaired in antisocial populations include dorsal and ventral regions of the prefrontal cortex (PFC), amygdala, hippocampus, angular gyrus, anterior cingulate and temporal cortex. Regions most commonly activated in moral judgment tasks consist of the polar/medial and ventral PFC, amygdala, angular gyrus and posterior cingulate. It is hypothesized that the rule-breaking behavior common to antisocial, violent and psychopathic individuals is in part due to impairments in some of the structures (dorsal and ventral PFC, amygdala and angular gyrus) subserving moral cognition and emotion. Impairments to the emotional component that comprises the feeling of what is moral is viewed as the primary deficit in antisocials, although some disruption to the cognitive and cognitive-emotional components of morality (particularly self-referential thinking and emotion regulation) cannot be ruled out. While this neurobiological predisposition is likely only one of several biosocial processes involved in the etiology of antisocial behavior, it raises significant moral issues for the legal system and neuroethics.
doi:10.1093/scan/nsl033
PMCID: PMC2555414  PMID: 18985107
antisocial; psychopathy; moral; prefrontal; temporal
15.  Satisfaction With Provider Communication Among Spanish-Speaking Medicaid Enrollees 
Objective
To determine if differences between English- and Spanish-speaking parents in ratings of their children’s health care can be explained by need for interpretive services.
Methods
Using the Consumer Assessment of Health Plans Survey–Child-Survey (CAHPS), reports about provider communication were compared among 3 groups of parents enrolled in a Medicaid managed care health plan: 1) English speakers, 2) Spanish speakers with no self-reported need for interpretive services, and 3) Spanish speakers with self-reported need for interpretive services. Parents were asked to report how well their providers 1) listened carefully to what was being said, 2) explained things in a way that could be understood, 3) respected their comments and concerns, and 4) spent enough time during medical encounters. Multivariate logistic regression was used to compare the ratings of each of the 3 groups while controlling for child’s gender, parent’s gender, parent’s educational attainment, child’s health status, and survey year.
Results
Spanish-speaking parents in need of interpretive services were less likely to report that providers spent enough time with their children (odds ratio = 0.34, 95% confidence interval = 0.17–0.68) compared to English-speaking parents. There was no statistically significant difference found between Spanish-speaking parents with no need of interpretive services and English-speaking parents.
Conclusions
Among Spanish- versus English-speaking parents, differences in ratings of whether providers spent enough time with children during medical encounters appear to be explained, in part, by need for interpretive services. No other differences in ratings of provider communication were found.
doi:10.1367/A04-019R1.1
PMCID: PMC1851901  PMID: 15548101
Consumer Assessment of Health Plan Survey; health disparities; Medicaid managed care; ratings of provider communication
16.  A Multisensory Cortical Network for Understanding Speech in Noise 
Journal of cognitive neuroscience  2009;21(9):1790-1805.
In noisy environments, listeners tend to hear a speaker’s voice yet struggle to understand what is said. The most effective way to improve intelligibility in such conditions is to watch the speaker’s mouth movements. Here we identify the neural networks that distinguish understanding from merely hearing speech, and determine how the brain applies visual information to improve intelligibility. Using functional magnetic resonance imaging, we show that understanding speech-in-noise is supported by a network of brain areas including the left superior parietal lobule, the motor/premotor cortex, and the left anterior superior temporal sulcus (STS), a likely apex of the acoustic processing hierarchy. Multisensory integration likely improves comprehension through improved communication between the left temporal–occipital boundary, the left medial-temporal lobe, and the left STS. This demonstrates how the brain uses information from multiple modalities to improve speech comprehension in naturalistic, acoustically adverse conditions.
doi:10.1162/jocn.2009.21118
PMCID: PMC2833290  PMID: 18823249
17.  Co-speech gestures influence neural activity in brain regions associated with processing semantic information 
Human brain mapping  2009;30(11):3509-3526.
Everyday communication is accompanied by visual information from several sources, including co-speech gestures, which provide semantic information listeners use to help disambiguate the speaker’s message. Using fMRI, we examined how gestures influence neural activity in brain regions associated with processing semantic information. The BOLD response was recorded while participants listened to stories under three audiovisual conditions and one auditory-only (speech alone) condition. In the first audiovisual condition, the storyteller produced gestures that naturally accompany speech. In the second, she made semantically unrelated hand movements. In the third, she kept her hands still. In addition to inferior parietal and posterior superior and middle temporal regions, bilateral posterior superior temporal sulcus and left anterior inferior frontal gyrus responded more strongly to speech when it was further accompanied by gesture, regardless of the semantic relation to speech. However, the right inferior frontal gyrus was sensitive to the semantic import of the hand movements, demonstrating more activity when hand movements were semantically unrelated to the accompanying speech. These findings show that perceiving hand movements during speech modulates the distributed pattern of neural activation involved in both biological motion perception and discourse comprehension, suggesting listeners attempt to find meaning, not only in the words speakers produce, but also in the hand movements that accompany speech.
doi:10.1002/hbm.20774
PMCID: PMC2896896  PMID: 19384890
discourse comprehension; fMRI; gestures; semantic processing; inferior frontal gyrus
18.  The amygdala and the experience of affect 
The current study examined the hypothesis that amygdala activation serves as a neural precondition for negative affective experience. Participants’ affective experience was measured by asking them to report on their momentary experiences several times a day over the course of a month using an electronic experience-sampling procedure. One year later, participants viewed backwardly masked depictions of fear while functional magnetic resonance imaging was used to measure their amygdala and fusiform gyrus activation. Negative affect, as measured during the experience-sampling procedure 1-year prior, was positively correlated with amygdala activation in response to these brief presentations of fear depictions. Furthermore, descriptive analyses indicated that fusiform gyrus activation and negative affective experience in the scanner were associated for participants reporting increased nervousness during the imaging procedure. The results are consistent with the interpretation that the amygdala contributes to negative affective experience by increasing perceptual sensitivity for negative stimuli.
doi:10.1093/scan/nsl042
PMCID: PMC2288526  PMID: 18392107
19.  The amygdala and the experience of affect 
The current study examined the hypothesis that amygdala activation serves as a neural precondition for negative affective experience. Participants’ affective experience was measured by asking them to report on their momentary experiences several times a day over the course of a month using an electronic experience-sampling procedure. One year later, participants viewed backwardly masked depictions of fear while functional magnetic resonance imaging was used to measure their amygdala and fusiform gyrus activation. Negative affect, as measured during the experience-sampling procedure 1-year prior, was positively correlated with amygdala activation in response to these brief presentations of fear depictions. Furthermore, descriptive analyses indicated that fusiform gyrus activation and negative affective experience in the scanner were associated for participants reporting increased nervousness during the imaging procedure. The results are consistent with the interpretation that the amygdala contributes to negative affective experience by increasing perceptual sensitivity for negative stimuli.
doi:10.1093/scan/nsl042
PMCID: PMC2288526  PMID: 18392107
20.  Brainstem Correlates of Speech-in-Noise Perception in Children 
Hearing research  2010;270(1-2):151-157.
Children often have difficulty understanding speech in challenging listening environments. In the absence of peripheral hearing loss, these speech perception difficulties may arise from dysfunction at more central levels in the auditory system, including subcortical structures. We examined brainstem encoding of pitch in a speech syllable in 38 school-age children. In children with poor speech-in-noise perception, we find impaired encoding of the fundamental frequency and the second harmonic, two important cues for pitch perception. Pitch, an important factor in speaker identification, aids the listener in tracking a specific voice from a background of voices. These results suggest that the robustness of subcortical neural encoding of pitch features in time-varying signals is an important factor in determining success with speech perception in noise.
doi:10.1016/j.heares.2010.08.001
PMCID: PMC2997182  PMID: 20708671
speech-in-noise perception; brainstem encoding; fundamental frequency; harmonics; noise exclusion; pitch
21.  Differential neural circuitry and self-interest in real vs hypothetical moral decisions 
Classic social psychology studies demonstrate that people can behave in ways that contradict their intentions—especially within the moral domain. We measured brain activity while subjects decided between financial self-benefit (earning money) and preventing physical harm (applying an electric shock) to a confederate under both real and hypothetical conditions. We found a shared neural network associated with empathic concern for both types of decisions. However, hypothetical and real moral decisions also recruited distinct neural circuitry: hypothetical moral decisions mapped closely onto the imagination network, while real moral decisions elicited activity in the bilateral amygdala and anterior cingulate—areas essential for social and affective processes. Moreover, during real moral decision-making, distinct regions of the prefrontal cortex (PFC) determined whether subjects make selfish or pro-social moral choices. Together, these results reveal not only differential neural mechanisms for real and hypothetical moral decisions but also that the nature of real moral decisions can be predicted by dissociable networks within the PFC.
doi:10.1093/scan/nss069
PMCID: PMC3475363  PMID: 22711879
real moral decision-making; fMRI; amygdala; TPJ; ACC
22.  Perception of Speech in Noise: Neural Correlates 
Journal of Cognitive Neuroscience  2010;23(9):2268-2279.
The presence of irrelevant auditory information (other talkers, environmental noises) presents a major challenge to listening to speech. The fundamental frequency (F0) of the target speaker is thought to provide an important cue for the extraction of the speaker’s voice from background noise, but little is known about the relationship between speech-in-noise (SIN) perceptual ability and neural encoding of the F0. Motivated by recent findings that music and language experience enhance brainstem representation of sound, we examined the hypothesis that brainstem encoding of the F0 is diminished to a greater degree by background noise in people with poorer perceptual abilities in noise. To this end, we measured speech-evoked auditory brainstem responses to /da/ in quiet and two multi-talker babble conditions (two-talker and six-talker) in native English-speaking young adults who ranged in their ability to perceive and recall SIN. Listeners who were poorer performers on a standardized SIN measure demonstrated greater susceptibility to the degradative effects of noise on the neural encoding of the F0. Particularly diminished was their phase-locked activity to the fundamental frequency in the portion of the syllable known to be most vulnerable to perceptual disruption (i.e., the formant transition period). Our findings suggest that the subcortical representation of the F0 in noise contributes to the perception of speech in noisy conditions.
doi:10.1162/jocn.2010.21556
PMCID: PMC3253852  PMID: 20681749
23.  Segmentation Cues in Conversational Speech: Robust Semantics and Fragile Phonotactics 
Multiple cues influence listeners’ segmentation of connected speech into words, but most previous studies have used stimuli elicited in careful readings rather than natural conversation. Discerning word boundaries in conversational speech may differ from the laboratory setting. In particular, a speaker’s articulatory effort – hyperarticulation vs. hypoarticulation (H&H) – may vary according to communicative demands, suggesting a compensatory relationship whereby acoustic-phonetic cues are attenuated when other information sources strongly guide segmentation. We examined how listeners’ interpretation of segmentation cues is affected by speech style (spontaneous conversation vs. read), using cross-modal identity priming. To elicit spontaneous stimuli, we used a map task in which speakers discussed routes around stylized landmarks. These landmarks were two-word phrases in which the strength of potential segmentation cues – semantic likelihood and cross-boundary diphone phonotactics – was systematically varied. Landmark-carrying utterances were transcribed and later re-recorded as read speech. Independent of speech style, we found an interaction between cue valence (favorable/unfavorable) and cue type (phonotactics/semantics). Thus, there was an effect of semantic plausibility, but no effect of cross-boundary phonotactics, indicating that the importance of phonotactic segmentation may have been overstated in studies where lexical information was artificially suppressed. These patterns were unaffected by whether the stimuli were elicited in a spontaneous or read context, even though the difference in speech styles was evident in a main effect. Durational analyses suggested speaker-driven cue trade-offs congruent with an H&H account, but these modulations did not impact on listener behavior. We conclude that previous research exploiting read speech is reliable in indicating the primacy of lexically based cues in the segmentation of natural conversational speech.
doi:10.3389/fpsyg.2012.00375
PMCID: PMC3464055  PMID: 23060839
speech segmentation; semantics; phonotactics; conversational speech; cross-modal priming
24.  Speaker Invariance for Phonetic Information: an fMRI Investigation 
Language and cognitive processes  2011;27(2):210-230.
The current study explored how listeners map the variable acoustic input onto a common sound structure representation while being able to retain phonetic detail to distinguish among the identity of talkers. An adaptation paradigm was utilized to examine areas which showed an equal neural response (equal release from adaptation) to phonetic change when spoken by the same speaker and when spoken by two different speakers, and insensitivity (failure to show release from adaptation) when the same phonetic input was spoken by a different speaker. Neural areas which showed speaker invariance were located in the anterior portion of the middle superior temporal gyrus bilaterally. These findings provide support for the view that speaker normalization processes allow for the translation of a variable speech input to a common abstract sound structure. That this process appears to occur early in the processing stream, recruiting temporal structures, suggests that this mapping takes place prelexically, before sound structure input is mapped on to lexical representations.
doi:10.1080/01690965.2011.594372
PMCID: PMC3528078  PMID: 23264714
25.  Cultural and linguistic effects on neural bases of ‘Theory of Mind’ in American and Japanese children 
Brain research  2007;1164:95-107.
Theory of mind (ToM) has been defined as our ability to predict behaviors of others in terms of their underlying intentions. While the developmental trajectory of ToM had been thought to be invariant across cultures, several ToM studies conducted outside the Anglo-American cultural or linguistic milieus have obtained mixed results. To examine effects of culture/language on the development of neural bases of ToM, we studied 12 American monolingual children and 12 Japanese bilingual children with second-order false-belief story and cartoon tasks, using functional magnetic resonance imaging (fMRI). While a few brain regions such as ventro-medial prefrontal cortex (vmPFC) and precuneus were recruited by the both cultural/linguistic groups, several brain areas including inferior frontal gyrus (IFG) and temporo-parietal junction (TPJ) were employed in a culture/language-dependent manner during the ToM tasks. These results suggest that the neural correlates of ToM may begin to vary depending upon cultural/linguistic background from early in life.
doi:10.1016/j.brainres.2007.06.022
PMCID: PMC2964053  PMID: 17643400

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