ARTICLE UPDATE - Functional dissociation of amygdala-modulated arousal and cognitive appraisal, in Turner syndrome

D. H. Skuse, J. S. Morris and R. J. Dolan

Brain, 128, 2084-2096

The amygdala is preferentially activated by facial expressions of fear. Right and left amygdala are hypothesized to play distinct, but complementary, roles that influence somatic and cognitive responses to facial expressions. Right amygdala activation is linked to autonomic arousal, and thus indirectly influences left hemisphere cognitive processing centres. Left amygdala activation is more closely associated with cognitive processing and differentiation of facial emotions. A double-dissociation between the functions of left and right amygdala is implied by lesion studies but supportive evidence is inconsistent, partly because patients with structural anteromedial temporal anomalies have experienced variable surgical procedures. A functional dissociation can be demonstrated between arousal and the cognitive appraisal of fearful faces in the condition of X-monosomy or Turner syndrome. Previous research found Turner syndrome women of normal verbal intelligence are seriously impaired in their ability cognitively to differentiate fearful from other facial expressions but they acquire fear conditioning normally, with enhanced autonomic responses. These findings supported the dissociation hypothesis, which was formally tested in a study of 12 X-monosomic and 12 control females who participated in functional magnetic resonance imaging during which simultaneous skin conductance recordings were acquired. Faces depicting fear or neutral emotions were presented to both case and control subjects in random order. Arousal to (fearful–neutral) faces was associated with transiently increased skin conductance responses and bilateral amygdala activation in both groups, but X-monosomic females had proportionately greater—and more persistent—right amygdala activation than controls. In both groups, cognitive accuracy correlated positively with differential activity of left fusiform gyrus. There was a significant correlation between the left fusiform and left medial amygdala activation only in normal females, and only in them did differential SCRs (to fearful–neutral faces) correlate positively with left fusiform responses. Arousal and cognitive appraisal functions of the amygdala can thus be functionally dissociated. X-monosomy selectively impairs explicit recognition of fearful faces in the presence of normal or enhanced autonomic reactivity, and is associated with a functional dissociation of activity in left amygdala and left fusiform gyrus. These findings imply X-linked genes are essential for binding somatic responses to the cognitive appraisal of emotional stimuli.

ARTICLE UPDATE - Neural evidence of effects of emotional valence on word recognition

Midori Inaba, Michio Nomura and Hideki Ohira

International Journal of Psychophysiology, 57, 165-173

There are no clear reports of electrophysiological evidence of the facilitating effect of negative valence on word recognition. However, behavioral psychological studies have suggested that negative words can be recognized more accurately than positive and neutral words. This study aimed to examine whether, and if so how, the valence of words could influence accuracy and event-related potentials (ERPs) in a recognition task. ERPs were recorded from 20 healthy subjects during performance of a word recognition task. We found a behavioral advantage in discriminability between old and new items for negative words. As for ERPs, the positive-going shift was evident for correct responses to targets in late latency at midline and left centro-parietal sites. Additionally, the magnitude of this component was greatest for negative targets, next for positive targets, and least for neutral targets. The findings offer support for the idea that negative content greatly accelerates recognition memory compared to positive and neutral words.

ARTICLE UPDATE - Memory for specific visual details can be enhanced by negative arousing content

Elizabeth A. Kensinger, Rachel J. Garoff-Eaton and Daniel L. Schacter

Journal of Memory and Language, 54(1), 99-112

Individuals often claim that they vividly remember information with negative emotional content. At least two types of information could lead to this sense of enhanced vividness: Information about the emotional item itself (e.g., the exact visual details of a snake) and information about the context in which the emotional item was encountered (e.g., the fact that the snake was sitting on a branch in a forest). The present study focused on the former, investigating how exposure duration at study and emotional content of an object affected the likelihood of remembering an item’s specific visual details. Participants studied neutral objects (e.g., a barometer) and negative arousing objects (e.g., a grenade) and were later shown either the identical object or a different photo of the same type of object (e.g., another barometer). Across two experiments, emotional content enhanced the likelihood that specific visual details were remembered: Individuals were more likely to correctly indicate that an item was identical to the object studied earlier if it was an emotional object than if it was a neutral object. This memory benefit for the emotional items was most robust when items were shown for longer exposure durations (500 or 1000 ms) rather than only briefly (for 250 ms). Thus, with sufficient processing time, negative arousing content appears to enhance the likelihood that visual details are remembered about an object.

ARTICLE UPDATE - Differential Effects of Lesions of the Amygdala and Prefrontal Cortex on Recognizing Facial Expressions of Complex Emotions

P. Shaw, J. Bramham, E. J. Lawrence, R. Morris, S. Baron-Cohen and A. S. David

Journal of Cognitive Neuroscience, 17, 1410-1419.

Humans can detect facial expressions of both simple, basic emotions and expressions reflecting more complex states of mind. The latter includes emotional expressions that regulate social interactions ("social expressions" such as looking hostile or friendly) and expressions that reflect the inner thought state of others ("cognitive expressions" such as looking pensive). To explore the neural substrate of this skill, we examined performance on a test of detection of such complex expressions in patients with lesions of the temporal lobe (n = 54) or frontal lobe (n = 31). Of the temporal group, 18 had unilateral focal lesions of the amygdala and of the frontal group, 14 patients had unilateral lesions of the ventromedial prefrontal cortex—two regions held to be pivotal in mediating social cognitive skills. Damage to either the left or right amygdala was associated with impairment in the recognition of both social and cognitive expressions, despite an intact ability to extract information relating to invariant physical attributes. Lesions to all of the right prefrontal cortex—not just the ventromedial portions—led to a specific deficit in recognizing complex social expressions with a negative valence. The deficit in the group with right prefrontal cortical damage may contribute to the disturbances in social behavior associated with such lesions. The results also suggest that the amygdala has a role in processing a wide range of emotional expressions.

ARTICLE UPDATE - Unattended Facial Expressions Asymmetrically Bias the Concurrent Processing of Nonemotional Information

Jeffrey S. Maxwell, Alexander J. Shackman and Richard J. Davidson

Journal of Cognitive Neuroscience, 17, 1386-1395.

Planned and reflexive behaviors often occur in the presence of emotional stimuli and within the context of an individual's acute emotional state. Therefore, determining the manner in which emotion and attention interact is an important step toward understanding how we function in the real world. Participants in the current investigation viewed centrally displayed, task-irrelevant, face distractors (angry, neutral, happy) while performing a lateralized go/no-go continuous performance task. Lateralized go targets and no-go lures that did not spatially overlap with the faces were employed to differentially probe processing in the left (LH) and right (RH) cerebral hemispheres. There was a significant interaction between expression and hemisphere, with an overall pattern such that angry distractors were associated with relatively more RH inhibitory errors than neutral or happy distractors and happy distractors with relatively more LH inhibitory errors than angry or neutral distractors. Simple effects analyses confirmed that angry faces differentially interfered with RH relative to LH inhibition and with inhibition in the RH relative to happy faces. A significant three-way interaction further revealed that state anxiety moderated relations between emotional expression and hemisphere. Under conditions of low cognitive load, more intense anxiety was associated with relatively greater RH than LH impairment in the presence of both happy and threatening distractors. By contrast, under high load, only angry distractors produced greater RH than LH interference as a function of anxiety.

ARTICLE UPDATE - The neurobiology of positive emotions

Burgdorf J, Panksepp J.

Neuroscience & Biobehavioral Reviews, in press

Compared to the study of negative emotions such as fear, the neurobiology of positive emotional processes and the associated positive affect (PA) states has only recently received scientific attention. Biological theories conceptualize PA as being related to (i) signals indicating that bodies are returning to equilibrium among those studying homeostasis, (ii) utility estimation among those favoring neuroeconomic views, and (iii) approach and other instinctual behaviors among those cultivating neuroethological perspectives. Indeed, there are probably several distinct forms of positive affect, but all are closely related to ancient sub-neocortical limbic brain regions we share with other mammals. There is now a convergence of evidence to suggest that various regions of the limbic system, including especially ventral striatal dopamine systems are implemented in an anticipatory (appetitive) positive affective state. Dopamine independent mechanisms utilizing opiate and GABA receptors in the ventral striatum, amygdala and orbital frontal cortex are important in elaborating consummatory PA (i.e. sensory pleasure) states, and various neuropeptides mediate homeostatic satisfactions.

ARTICLE UPDATE - On the role of attention for the processing of emotions in speech: Sex differences revisited.

Schirmer A, Kotz SA, Friederici AD.

Cognitive Brain Research, 24, 442-52.

In a previous cross-modal priming study [A. Schirmer, A.S. Kotz, A.D. Friederici, Sex differentiates the role of emotional prosody during word processing, Cogn. Brain Res. 14 (2002) 228-233.], we found that women integrated emotional prosody and word valence earlier than men. Both sexes showed a smaller N400 in the event-related potential to emotional words when these words were preceded by a sentence with congruous compared to incongruous emotional prosody. However, women showed this effect with a 200-ms interval between prime sentence and target word whereas men showed the effect with a 750-ms interval. The present study was designed to determine whether these sex differences prevail when attention is directed towards the emotional content of prosody and word meaning. To this end, we presented the same prime sentences and target words as in our previous study. Sentences were spoken with happy or sad prosody and followed by a congruous or incongruous emotional word or pseudoword. The interval between sentence offset and target onset was 200 ms. In addition to performing a lexical decision, participants were asked to decide whether or not a word matched the emotional prosody of the preceding sentence. The combined lexical and congruence judgment failed to reveal differences in emotional-prosodic priming between men and women. Both sexes showed smaller N400 amplitudes to emotionally congruent compared to incongruent words. This suggests that the presence of sex differences in emotional-prosodic priming depends on whether or not participants are instructed to take emotional prosody into account.

ARTICLE UPDATE - Incidental effects of emotional valence in single word processing: An fMRI study

Lars Kuchinke, Arthur M. Jacobs, Claudia Grubich, Melissa L.-H. Võ, Markus Conrada and Manfred Herrmann

NeuroImage, 28, 1022-1032

The present study aimed at identifying the neural responses associated with the incidental processing of the emotional valence of single words using event-related functional magnetic resonance imaging (fMRI). Twenty right-handed participants performed a visual lexical decision task, discriminating between nouns and orthographically and phonologically legal nonwords. Positive, neutral and negative word categories were matched for frequency, number and frequency of orthographic neighbors, number of letters and imageability. Response times and accuracy data differed significantly between positive and neutral, and positive and negative words respectively, thus, replicating the findings of a pilot study. Words showed distributed, mainly left hemisphere activations, indicating involvement of a neural network responsible for semantic word knowledge. The neuroimaging data further revealed areas in left orbitofrontal gyrus and bilateral inferior frontal gyrus with greater activation to emotional than to neutral words. These brain regions are known to be involved in processing semantic and emotional information. Furthermore, distinct activations associated with positive words were observed in bilateral middle temporal and superior frontal gyrus, known to support semantic retrieval, and a distributed network, namely anterior and posterior cingulate gyrus, lingual gyrus and hippocampus when comparing positive and negative words. The latter areas were previously associated with explicit and not incidental processing of the emotional meaning of words and emotional memory retrieval. Thus, the results are discussed in relation to models of processing semantic and episodic emotional information.

ARTICLE UPDATE - Functional consequences of perceiving facial expressions of emotion without awareness.

Eastwood JD, Smilek D.

Consciousness & Cognition, 14, 565-84

A substantial body of research has established that even when we are not consciously aware of the faces of others we are nevertheless sensitive to, and impacted by their facial expression. In this paper, we consider this body of research from a new perspective by examining the functions of unconscious perception revealed by these studies. A consideration of the literature from this perspective highlights that existing research methods are limited when it comes to revealing possible functions of unconscious perception. The critical shortcoming is that in all of the methods, the perceived facial expression remains outside of awareness. This is a problem because there are good reasons to believe that one important function of unconsciously perceived negative faces is to attract attention so that they are consciously perceived; such conscious perception, however, is never allowed with existing methodologies. We discuss recent studies of emotional face perception under conditions of visual search that address this issue directly. Further, we suggest that methodologies that do not examine cognitive processes as they occur in more natural settings may result in fundamental misunderstandings of human cognition.

ARTICLE UPDATE - How do we modulate our emotions? Parametric fMRI reveals cortical midline structures as regions specifically involved in the processi

Alexander Heinzel, Felix Bermpohl, Robert Niese, Andrea Pfennig, Alvaro Pascual-Leone, Gottfried Schlaug and Georg Northoff

Cognitive Brain Research, 25,348-358.

One of the major problems in affective neuroscience of healthy subjects as well as of patients with emotional dysfunctions is to disentangle emotional core functions and non-emotional processes. Emotional valence is considered an emotional key process. The present study employed a parametric functional magnetic resonance imaging (fMRI) study to address this question. Thirteen healthy volunteers were scanned during emotional stimulus processing (International Affective Picture System). The presented pictures covered the entire range of emotional valences. The fMRI data were consecutively subjected to a preliminary categorical (valence-independent) and a detailed parametric analysis, the latter using individual valence ratings as regressor. The parametric analysis revealed a linear valence-dependent modulation of the BOLD signal in the orbito- and dorsomedial prefrontal cortex (OMPFC, DMPFC), medial parietal cortex (MPC), and insula. In addition, we observed that emotional valence exerts its effects predominantly via modulation of signal decreases. We conclude that the psychological concept of emotional valence may be related to neural processing in cortical midline regions.

ARTICLE UPDATE - Signals for threat modulate attentional capture and holding: Fear‐conditioning and extinction during the exogenous cueing task

Ernst H. W. Koster , Geert Crombez , Stefaan Van Damme , Bruno Verschuere , Jan De Houwer

Cognition & Emotion, 18, 771-780.

This experiment investigated whether acquired signals for threat capture and hold visual attention. Cues that were presented in an exogenous attentional cueing task were emotionally modulated using a fear‐conditioning paradigm. During acquisition, undergraduate students (55 women, 11 men) learned that one cue (CS+) of the attentional task was a signal for an aversive white noise burst (UCS, 100 dBA) and that another cue (CS−) signalled its nonoccurrence. In a subsequent extinction phase, no UCSs were presented anymore during the cueing task. Results indicated that during acquisition, the CS+ cues strongly captured and held visual attention in comparison with the CS− cues, and that these effects diminished during extinction.

ARTICLE UPDATE - Affective priming as an indirect measure of food preferences acquired through odor conditioning.

Hermans D, Baeyens F, Lamote S, Spruyt A, Eelen P.

Experimental Psychology,52, 180-186.

The present study aimed at investigating affective priming for originally neutral food stimuli that recently acquired their affective meaning through odor conditioning. In a first phase, pictures of different brands of yogurts (CSs) were contingently presented with a positive or negative odor (US). In a subsequent phase, the yogurt CSs were used as primes in an affective priming procedure. Rating data showed that the acquisition procedure resulted in a reliable evaluative learning effect. This could be corroborated by the results of the priming task. Participants responded faster to positive target words and made fewer errors when they were preceded by a CS that had been associated with a positive odor, as compared to a CS that was associated with a negative odor. A reversed pattern was present for negative targets. Based on these findings, it is suggested that affective priming might be used as a demand-free measure of evaluative learning.

ARTICLE UPDATE - When less is more: the consequences of affective primacy for subliminal priming effects.

Stapel DA, Koomen W.

Personality & Socical Psychological Bulletin, 31, 1286-1295.

This research investigates the consequences of the notion that one can distinguish early-evaluative (when exposure is short) and late-descriptive reactions (when exposure is long) to subliminally primed trait concepts. In three studies, it was found that the evaluative effects instigated by short exposure to primed concepts were bigger than the evaluative + descriptive effects instigated by long exposure: Less is more. Only when exposure was short, target interpretations were accompanied by evaluative inferences (Studies 1 and 3). Similarly, only when exposure was short, descriptively inapplicable trait primes affected the interpretation of an ambiguous target (Studies 2 and 3).

ARTICLE UPDATE - Cognitive-emotional distinctiveness: Separating emotions from non-emotions in the representation of a stressful memory.

Boals A, Klein K.

Memory, 13,638-648.

Current theories on autobiographical memory and recent neurological evidence suggest that emotional and non-emotional features of a memory may be retrieved by separate systems. To test this notion, 207 participants who had experienced the break-up of a significant romantic relationship in the last 12 months completed a Multidimensional Scaling (MDS) procedure in relation to the previous relationship. The resulting MDS model revealed two dimensions: a valence and an emotional/non-emotional dimension. Further, participants who associated a high level of distress with their relationship break-up perceived less dissimilarity between emotional and non-emotional features than participants who associated a low level of distress with their relationship break-up. Theoretical and methodological implications for stress and memory are discussed.

ARTICLE UPDATE - Investigating emotion with music: An fMRI study.

Koelsch S, Fritz T, V Cramon DY, Muller K, Friederici AD.

Human Brain Mapping, in press

The present study used pleasant and unpleasant music to evoke emotion and functional magnetic resonance imaging (fMRI) to determine neural correlates of emotion processing. Unpleasant (permanently dissonant) music contrasted with pleasant (consonant) music showed activations of amygdala, hippocampus, parahippocampal gyrus, and temporal poles. These structures have previously been implicated in the emotional processing of stimuli with (negative) emotional valence; the present data show that a cerebral network comprising these structures can be activated during the perception of auditory (musical) information. Pleasant (contrasted to unpleasant) music showed activations of the inferior frontal gyrus (IFG, inferior Brodmann's area (BA) 44, BA 45, and BA 46), the anterior superior insula, the ventral striatum, Heschl's gyrus, and the Rolandic operculum. IFG activations appear to reflect processes of music-syntactic analysis and working memory operations. Activations of Rolandic opercular areas possibly reflect the activation of mirror-function mechanisms during the perception of the pleasant tunes. Rolandic operculum, anterior superior insula, and ventral striatum may form a motor-related circuitry that serves the formation of (premotor) representations for vocal sound production during the perception of pleasant auditory information. In all of the mentioned structures, except the hippocampus, activations increased over time during the presentation of the musical stimuli, indicating that the effects of emotion processing have temporal dynamics; the temporal dynamics of emotion have so far mainly been neglected in the functional imaging literature.

ARTICLE UPDATE - Unintentional processing of motivational valence

Agnes Moors, Jan De Houwer, Dirk Hermans, Paul Eelen

The Quarterly Journal of Experimental Psychology: Section A, 58, 1043-1063

Recent motivational affective priming studies (Moors & De Houwer, 2001; Moors, De Houwer, & Eelen, 2004) showed that primes that indicate success on a goal-inducing task facilitate positive target responses whereas primes that indicate failure on that task facilitate negative target responses. In the current studies, we examined whether these priming effects depend on consciously intentional processing of the motivational valence of the primes. In Experiment 1, the outcome of success or failure was presented not only immediately before the target (i.e., the prime) but also a second time after the target response. This should encourage participants to ignore the prime. In Experiment 2, participants were asked to respond to the targets within 600 ms after target onset. As a result, participants had little opportunity to process the motivational prime valence in a consciously intentional way. Nevertheless, strong affective priming effects were found in both studies. These results provide additional support for the claim that motivational valence can be processed without the conscious intention to do so.

ARTICLE UPDATE - Emotion and attention interactions in social cognition: Brain regions involved in processing anger prosody

David Sander, Didier Grandjean, Gilles Pourtois, Sophie Schwartz, Mohamed L. Seghier, Klaus R. Scherer and Patrik Vuilleumier

NeuroImage, 28,848-858

Multiple levels of processing are thought to be involved in the appraisal of emotionally relevant events, with some processes being engaged relatively independently of attention, whereas other processes may depend on attention and current task goals or context. We conducted an event-related fMRI experiment to examine how processing angry voice prosody, an affectively and socially salient signal, is modulated by voluntary attention. To manipulate attention orthogonally to emotional prosody, we used a dichotic listening paradigm in which meaningless utterances, pronounced with either angry or neutral prosody, were presented simultaneously to both ears on each trial. In two successive blocks, participants selectively attended to either the left or right ear and performed a gender-decision on the voice heard on the target side. Our results revealed a functional dissociation between different brain areas. Whereas the right amygdala and bilateral superior temporal sulcus responded to anger prosody irrespective of whether it was heard from a to-be-attended or to-be-ignored voice, the orbitofrontal cortex and the cuneus in medial occipital cortex showed greater activation to the same emotional stimuli when the angry voice was to-be-attended rather than to-be-ignored. Furthermore, regression analyses revealed a strong correlation between orbitofrontal regions and sensitivity on a behavioral inhibition scale measuring proneness to anxiety reactions. Our results underscore the importance of emotion and attention interactions in social cognition by demonstrating that multiple levels of processing are involved in the appraisal of emotionally relevant cues in voices, and by showing a modulation of some emotional responses by both the current task-demands and individual differences.