John G. Taylor & Nickolaos F. Fragopanagos
Neural Networks, in press
We analyse emotions from the viewpoint of how emotion and attention interact in the brain. Much has been learnt about the brain structures involved in attention, especially in vision. In particular the manner in which attention functions as a high-level control system, able to make cognitive processing so effective, has been studied both at a global level by brain imaging (fMRI, PET, MEG and EEG), at a local single cell level in monkeys and lower animals, and computationally by a variety of models. The manner in which emotions impinge on this attention control system is not so well analysed, although numerous new results are now emerging from using the same tools. Here we use an engineering control approach to attention to model it in a global manner but with relatively sure local foundations at singe neuron level.
The manner in which emotional value (as coded in amygdale and orbito-frontal cortex) can interact with the attention control circuitry is analysed using results of various experimental paradigms. A general model of this interaction is first developed and tested against a list of paradigms, and then more detailed computations are performed using more specific features of the attention control system and the limbic value coding. These computations are completed by a simulation of the emotional attentional blink, a demanding paradigm for any model of attention alone, but made more so by the presence of emotional value codes for stimuli. We conclude the paper with a general discussion of further avenues of research.
Monday, May 30, 2005
ARTICLE UPDATE - Retrieving accurate and distorted memories: Neuroimaging evidence for effects of emotion
Elizabeth A. KensingeR & Daniel Schacter
NeuroImage, in press
While limbic activity is known to be associated with successful encoding of emotional information, it is less clear whether it is related to successful retrieval. The present fMRI study assessed the effects of emotion on the neural processes engaged during retrieval of accurate compared to distorted memories. Prior to the scan, participants (16 young adults) viewed names of neutral (e.g., frog) and emotional (e.g., snake) objects and formed a mental image of the object named. They were shown photos of half of the objects. During the fMRI scan, participants saw object names and indicated whether or not they had seen the corresponding photo. Memory distortions (misattributions) occurred when participants incorrectly indicated whether or not a photo had been studied. Activity in some regions (e.g., L anterior hippocampus) was related to accurate retrieval (correct attributions > misattributions) for emotional and neutral items. However, activity in other regions corresponded with accurate retrieval specifically for emotional items (e.g., in R amygdala/periamygdaloid cortex and L orbitofrontal cortex) or for neutral items (e.g., in lateral inferior prefrontal cortex and R posterior hippocampus). Results indicate that emotional salience modulates the processes engaged during accurate retrieval and that activity in limbic regions corresponds with accurate memory assignment for emotional items. To our knowledge, this study is the first to demonstrate a link between limbic engagement at retrieval and accurate memory attribution.
NeuroImage, in press
While limbic activity is known to be associated with successful encoding of emotional information, it is less clear whether it is related to successful retrieval. The present fMRI study assessed the effects of emotion on the neural processes engaged during retrieval of accurate compared to distorted memories. Prior to the scan, participants (16 young adults) viewed names of neutral (e.g., frog) and emotional (e.g., snake) objects and formed a mental image of the object named. They were shown photos of half of the objects. During the fMRI scan, participants saw object names and indicated whether or not they had seen the corresponding photo. Memory distortions (misattributions) occurred when participants incorrectly indicated whether or not a photo had been studied. Activity in some regions (e.g., L anterior hippocampus) was related to accurate retrieval (correct attributions > misattributions) for emotional and neutral items. However, activity in other regions corresponded with accurate retrieval specifically for emotional items (e.g., in R amygdala/periamygdaloid cortex and L orbitofrontal cortex) or for neutral items (e.g., in lateral inferior prefrontal cortex and R posterior hippocampus). Results indicate that emotional salience modulates the processes engaged during accurate retrieval and that activity in limbic regions corresponds with accurate memory assignment for emotional items. To our knowledge, this study is the first to demonstrate a link between limbic engagement at retrieval and accurate memory attribution.
Wednesday, May 18, 2005
ARTICLE UPDATE - The neural bases of amusement and sadness: A comparison of block contrast and subject-specific emotion intensity regression approache
Goldin PR, Hutcherson CA, Ochsner KN, Glover GH, Gabrieli JD, Gross JJ.
Neuroimage, in press
Neuroimaging studies have made substantial progress in elucidating the neural bases of emotion. However, few studies to date have directly addressed the subject-specific, time-varying nature of emotional responding. In the present study, we employed functional magnetic resonance imaging to examine the neural bases of two common emotions-amusement and sadness-using both (a) a stimulus-based block contrast approach and (b) a subject-specific regression analysis using continuous ratings of emotional intensity. Thirteen women viewed a set of nine 2-min amusing, sad, or neutral film clips two times. During the first viewing, participants watched the film stimuli. During the second viewing, they made continuous ratings of the intensity of their own amusement and sadness during the first film viewing. For sad films, both block contrast and subject-specific regression approaches resulted in activations in medial prefrontal cortex, inferior frontal gyrus, superior temporal gyrus, precuneus, lingual gyrus, amygdala, and thalamus. For amusing films, the subject-specific regression analysis demonstrated significant activations not detected by the block contrast in medial, inferior frontal gyrus, dorsolateral prefrontal cortex, posterior cingulate, temporal lobes, hippocampus, thalamus, and caudate. These results suggest a relationship between emotion-specific temporal dynamics and the sensitivity of different data analytic methods for identifying emotion-related neural responses. These findings shed light on the neural bases of amusement and sadness, and highlight the value of using emotional film stimuli and subject-specific continuous emotion ratings to characterize the dynamic, time-varying components of emotional responses.
Neuroimage, in press
Neuroimaging studies have made substantial progress in elucidating the neural bases of emotion. However, few studies to date have directly addressed the subject-specific, time-varying nature of emotional responding. In the present study, we employed functional magnetic resonance imaging to examine the neural bases of two common emotions-amusement and sadness-using both (a) a stimulus-based block contrast approach and (b) a subject-specific regression analysis using continuous ratings of emotional intensity. Thirteen women viewed a set of nine 2-min amusing, sad, or neutral film clips two times. During the first viewing, participants watched the film stimuli. During the second viewing, they made continuous ratings of the intensity of their own amusement and sadness during the first film viewing. For sad films, both block contrast and subject-specific regression approaches resulted in activations in medial prefrontal cortex, inferior frontal gyrus, superior temporal gyrus, precuneus, lingual gyrus, amygdala, and thalamus. For amusing films, the subject-specific regression analysis demonstrated significant activations not detected by the block contrast in medial, inferior frontal gyrus, dorsolateral prefrontal cortex, posterior cingulate, temporal lobes, hippocampus, thalamus, and caudate. These results suggest a relationship between emotion-specific temporal dynamics and the sensitivity of different data analytic methods for identifying emotion-related neural responses. These findings shed light on the neural bases of amusement and sadness, and highlight the value of using emotional film stimuli and subject-specific continuous emotion ratings to characterize the dynamic, time-varying components of emotional responses.
ARTICLE UPDATE - A direct intracranial record of emotions evoked by subliminal words
Lionel Naccache, Raphaël Gaillard, Claude Adam, Dominique Hasboun, Stéphane Clémenceau, Michel Baulac, Stanislas Dehaene, and Laurent Cohen
PNAS, 102, 7713-7717.
A classical but still open issue in cognitive psychology concerns the depth of subliminal processing. Can the meaning of undetected words be accessed in the absence of consciousness? Subliminal priming experiments in normal subjects have revealed only small effects whose interpretation remains controversial. Here, we provide a direct demonstration of semantic access for unseen masked words. In three epileptic patients with intracranial electrodes, we recorded brain potentials from the amygdala, a neural structure that responds to fearful or threatening stimuli presented in various modalities, including written words. We show that the subliminal presentation of emotional words modulates the activity of the amygdala at a long latency (>800 ms). Our result indicates that subliminal words can trigger long-lasting cerebral processes, including semantic access to emotional valence.
PNAS, 102, 7713-7717.
A classical but still open issue in cognitive psychology concerns the depth of subliminal processing. Can the meaning of undetected words be accessed in the absence of consciousness? Subliminal priming experiments in normal subjects have revealed only small effects whose interpretation remains controversial. Here, we provide a direct demonstration of semantic access for unseen masked words. In three epileptic patients with intracranial electrodes, we recorded brain potentials from the amygdala, a neural structure that responds to fearful or threatening stimuli presented in various modalities, including written words. We show that the subliminal presentation of emotional words modulates the activity of the amygdala at a long latency (>800 ms). Our result indicates that subliminal words can trigger long-lasting cerebral processes, including semantic access to emotional valence.
Tuesday, May 17, 2005
ARTICLE UPDATE - Binocular rivalry between emotional and neutral stimuli: A validation using fear conditioning and EEG
Georg W. Alpers, Mirjana Ruhleder, Nora Walz, Andreas Mühlberger and Paul Pauli
International Journal of Psychophysiology, in press
When two incompatible pictures are projected to the two eyes, they compete for perceptual dominance. Previous research has claimed that meaningful and emotionally valenced pictures predominate over neutral pictures in this rivalry. This may be interpreted as evidence for preferential processing of emotionally significant stimuli in the visual system but it is difficult to dismiss that the physical characteristics of the different pictures or response biases influenced the results of these studies. Thus, we set out to examine the influence of emotion using methods eliminating the influence of physical characteristics and minimizing response biases. We used simple visual patterns and induced emotional valence by fear conditioning. In Experiment 1 the aversive CS+ predominated over the CS−. In Experiment 2 we extended previous findings by showing that participants' self-reported perception is validated by corresponding steady-state visually evoked potentials in the EEG in the context of such a conditioning experiment. This was accomplished by frequency coding the rivalling stimuli with a stimulus specific pattern reversal and extracting the corresponding frequency from the occipital lobe EEG. Taken together, these studies provide further evidence that picture valence can influence perception in binocular rivalry. This is discussed in terms of subcortical mechanisms supporting the efficient processing of threatening information.
International Journal of Psychophysiology, in press
When two incompatible pictures are projected to the two eyes, they compete for perceptual dominance. Previous research has claimed that meaningful and emotionally valenced pictures predominate over neutral pictures in this rivalry. This may be interpreted as evidence for preferential processing of emotionally significant stimuli in the visual system but it is difficult to dismiss that the physical characteristics of the different pictures or response biases influenced the results of these studies. Thus, we set out to examine the influence of emotion using methods eliminating the influence of physical characteristics and minimizing response biases. We used simple visual patterns and induced emotional valence by fear conditioning. In Experiment 1 the aversive CS+ predominated over the CS−. In Experiment 2 we extended previous findings by showing that participants' self-reported perception is validated by corresponding steady-state visually evoked potentials in the EEG in the context of such a conditioning experiment. This was accomplished by frequency coding the rivalling stimuli with a stimulus specific pattern reversal and extracting the corresponding frequency from the occipital lobe EEG. Taken together, these studies provide further evidence that picture valence can influence perception in binocular rivalry. This is discussed in terms of subcortical mechanisms supporting the efficient processing of threatening information.
Tuesday, May 10, 2005
ARTICLE UPDATE - Functional connectivity with anterior cingulate and orbitofrontal cortices during decision-making
M.X Cohen, A.S. Heller and C. Ranganath
Cognitive Brain Research, 23, 61-70
Recent neuroscience research is beginning to discover the brain regions involved in decision-making under uncertainty, but little is known about whether or how these regions functionally interact with each other. Here, we used event-related functional magnetic resonance imaging to examine both changes in overall activity and changes in functional connectivity during risk-taking. Results showed that choosing high-risk over low-risk decisions was associated with increased activity in both anterior cingulate and orbitofrontal cortices. Connectivity analyses revealed that largely distinct, but somewhat overlapping, cortical and subcortical regions exhibited significant functional connectivity with anterior cingulate and orbitofrontal cortices. Additionally, connectivity with the anterior cingulate in some regions, including the orbitofrontal cortex and nucleus accumbens, was modulated by the decision participants chose. These findings (1) elucidate large networks of brain regions that are functionally connected with both anterior cingulate and orbitofrontal cortices during decision-making and (2) demonstrate that the roles of orbitofrontal and anterior cingulate cortices can be functionally differentiated by examining patterns of connectivity.
Cognitive Brain Research, 23, 61-70
Recent neuroscience research is beginning to discover the brain regions involved in decision-making under uncertainty, but little is known about whether or how these regions functionally interact with each other. Here, we used event-related functional magnetic resonance imaging to examine both changes in overall activity and changes in functional connectivity during risk-taking. Results showed that choosing high-risk over low-risk decisions was associated with increased activity in both anterior cingulate and orbitofrontal cortices. Connectivity analyses revealed that largely distinct, but somewhat overlapping, cortical and subcortical regions exhibited significant functional connectivity with anterior cingulate and orbitofrontal cortices. Additionally, connectivity with the anterior cingulate in some regions, including the orbitofrontal cortex and nucleus accumbens, was modulated by the decision participants chose. These findings (1) elucidate large networks of brain regions that are functionally connected with both anterior cingulate and orbitofrontal cortices during decision-making and (2) demonstrate that the roles of orbitofrontal and anterior cingulate cortices can be functionally differentiated by examining patterns of connectivity.
Thursday, May 05, 2005
ARTICLE UPDATE - The cognitive control of emotion
Kevin N. Ochsner and James J. Gross
Trends in Cognitive Sciences, 9, 242-249
The capacity to control emotion is important for human adaptation. Questions about the neural bases of emotion regulation have recently taken on new importance, as functional imaging studies in humans have permitted direct investigation of control strategies that draw upon higher cognitive processes difficult to study in nonhumans. Such studies have examined (1) controlling attention to, and (2) cognitively changing the meaning of, emotionally evocative stimuli. These two forms of emotion regulation depend upon interactions between prefrontal and cingulate control systems and cortical and subcortical emotion-generative systems. Taken together, the results suggest a functional architecture for the cognitive control of emotion that dovetails with findings from other human and nonhuman research on emotion.
Trends in Cognitive Sciences, 9, 242-249
The capacity to control emotion is important for human adaptation. Questions about the neural bases of emotion regulation have recently taken on new importance, as functional imaging studies in humans have permitted direct investigation of control strategies that draw upon higher cognitive processes difficult to study in nonhumans. Such studies have examined (1) controlling attention to, and (2) cognitively changing the meaning of, emotionally evocative stimuli. These two forms of emotion regulation depend upon interactions between prefrontal and cingulate control systems and cortical and subcortical emotion-generative systems. Taken together, the results suggest a functional architecture for the cognitive control of emotion that dovetails with findings from other human and nonhuman research on emotion.
ARTICLE UPDATE - Same or different? Neural correlates of happy and sad mood in healthy males
Ute Habel, Martina Klein, Thilo Kellermann, N. Jon Shah and Frank Schneider
NeuroImage, 26, 206-214
Emotional experience in healthy men has been shown to rely on a brain network including subcortical as well as cortical areas in a complex interaction, which may be substantially influenced by many internal personal and external factors such as individuality, gender, stimulus material and task instructions. The divergent results may be interpreted by taking these considerations into account. Hence, many aspects remain to be clarified in characterizing the neural correlates underlying the subjective experience of emotion. One unresolved question refers to the influence of emotion quality on the cerebral substrates. Hence, 26 male healthy subjects were investigated with functional magnetic resonance imaging during standardized sad and happy mood induction as well as a cognitive control task to explore brain responses differentially involved in positive and negative emotional experience. Sad and happy mood in contrast to the control task produced similarly significant activations in the amygdala–hippocampal area extending into the parahippocampal gyrus as well as in the prefrontal and temporal cortex, the anterior cingulate, and the precuneus. Significant valence differences emerged when comparing both tasks directly. More activation has been demonstrated in the ventrolateral prefrontal cortex (VLPFC), the anterior cingulate cortex (ACC), the transverse temporal gyrus, and the superior temporal gyrus during sadness. Happiness, on the other hand, produced stronger activations in the dorsolateral prefrontal cortex (DLPFC), the cingulate gyrus, the inferior temporal gyrus, and the cerebellum. Hence, negative and positive moods reveal distinct cortical activation foci within a common neural network, probably making the difference between qualitatively different emotional feelings.
NeuroImage, 26, 206-214
Emotional experience in healthy men has been shown to rely on a brain network including subcortical as well as cortical areas in a complex interaction, which may be substantially influenced by many internal personal and external factors such as individuality, gender, stimulus material and task instructions. The divergent results may be interpreted by taking these considerations into account. Hence, many aspects remain to be clarified in characterizing the neural correlates underlying the subjective experience of emotion. One unresolved question refers to the influence of emotion quality on the cerebral substrates. Hence, 26 male healthy subjects were investigated with functional magnetic resonance imaging during standardized sad and happy mood induction as well as a cognitive control task to explore brain responses differentially involved in positive and negative emotional experience. Sad and happy mood in contrast to the control task produced similarly significant activations in the amygdala–hippocampal area extending into the parahippocampal gyrus as well as in the prefrontal and temporal cortex, the anterior cingulate, and the precuneus. Significant valence differences emerged when comparing both tasks directly. More activation has been demonstrated in the ventrolateral prefrontal cortex (VLPFC), the anterior cingulate cortex (ACC), the transverse temporal gyrus, and the superior temporal gyrus during sadness. Happiness, on the other hand, produced stronger activations in the dorsolateral prefrontal cortex (DLPFC), the cingulate gyrus, the inferior temporal gyrus, and the cerebellum. Hence, negative and positive moods reveal distinct cortical activation foci within a common neural network, probably making the difference between qualitatively different emotional feelings.
ARTICLE UPDATE - Activity and Connectivity of Brain Mood Regulating Circuit in Depression: A Functional Magnetic Resonance Study
Amit Anand, Yu Li, Yang Wang, Jingwei Wu, Sujuan Gao, Lubna Bukharia, Vincent P. Mathews, Andrew Kalnin and Mark J. Lowe
Biological Psychiatry, 57, 1079-1088
Background
Functional imaging studies indicate that imbalances in cortico-limbic activity and connectivity may underlie the pathophysiology of MDD. In this study, using functional Magnetic Resonance Imaging (fMRI), we investigated differences in cortico-limbic activity and connectivity between depressed patients and healthy controls.
Methods
Fifteen unmedicated unipolar depressed patients and 15 matched healthy subjects underwent fMRI during which they first completed a conventional block-design activation experiment in which they were exposed to negative and neutral pictures. Next, low frequency blood oxygenation dependent (BOLD) related fluctuations (LFBF) data were acquired at rest and during steady-state exposure to neutral, positive and negative pictures. LFBF correlations were calculated between anterior cingulate cortex (ACC) and limbic regions – amygdala (AMYG), pallidostriatum (PST) and medial thalamus (MTHAL) and used as a measure of cortico-limbic connectivity.
Results
Depressed patients had increased activation of cortical and limbic regions. At rest and during exposure to neutral, positive, and negative pictures cortico-limbic LFBF correlations were decreased in depressed patients compared to healthy subjects.
Conclusions
The finding of increased activation of limbic regions and decreased LFBF correlations between ACC and limbic regions is consistent with the hypothesis that decreased cortical regulation of limbic activation in response to negative stimuli may be present in depression.
Biological Psychiatry, 57, 1079-1088
Background
Functional imaging studies indicate that imbalances in cortico-limbic activity and connectivity may underlie the pathophysiology of MDD. In this study, using functional Magnetic Resonance Imaging (fMRI), we investigated differences in cortico-limbic activity and connectivity between depressed patients and healthy controls.
Methods
Fifteen unmedicated unipolar depressed patients and 15 matched healthy subjects underwent fMRI during which they first completed a conventional block-design activation experiment in which they were exposed to negative and neutral pictures. Next, low frequency blood oxygenation dependent (BOLD) related fluctuations (LFBF) data were acquired at rest and during steady-state exposure to neutral, positive and negative pictures. LFBF correlations were calculated between anterior cingulate cortex (ACC) and limbic regions – amygdala (AMYG), pallidostriatum (PST) and medial thalamus (MTHAL) and used as a measure of cortico-limbic connectivity.
Results
Depressed patients had increased activation of cortical and limbic regions. At rest and during exposure to neutral, positive, and negative pictures cortico-limbic LFBF correlations were decreased in depressed patients compared to healthy subjects.
Conclusions
The finding of increased activation of limbic regions and decreased LFBF correlations between ACC and limbic regions is consistent with the hypothesis that decreased cortical regulation of limbic activation in response to negative stimuli may be present in depression.
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