CEREBRO Y EMOCIO N

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Hartikainen KM, Ogawa KH, Soltani M, Knight RT

Emotionally arousing stimuli compete for attention with left hemispace.
Neuroreport. 2007 Dec 3;18(18):1929-33.

Rapid interaction of the emotional and attentional networks is critical for adaptive
behavior. Here, we examined the effects of emotional stimulation on hemifield
attention allocation using event-related potential and behavioral measures.
Participants performed a visual-discrimination task on nonemotional targets
presented randomly in the left or right hemifield. A brief task-irrelevant emotional
(pleasant or unpleasant; 150-ms duration) or neutral picture was presented
centrally 350 ms before the next target (150-ms duration). Unpleasant stimuli
interfered with the left visual field attention capacity, slowing behavioral
responses to attended left field stimuli. In keeping with the behavioral data,
event-related potential responses to nonemotional attended left field stimuli were
reduced over the right parietal regions when preceded by an unpleasant event.
The results provide electrophysiological and behavioral evidence that
unpleasant, emotionally arousing stimuli interfere with the right hemisphere-
dependent attention capacity.

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Shamay-Tsoory SG, Tibi-Elhanany Y, Aharon-Peretz J The green-eyed

monster and malicious joy: the neuroanatomical bases of envy and gloating
(schadenfreude). Brain. 2007 Jun;130(Pt 6):1663-78
.

Facing a protagonist's emotional mental state can trigger social emotions (or
'fortune of others' emotion), such as envy or gloating, which reflect one's
assessment of the consequences of the other's fortune. Here we suggest that
these social emotions are mediated by the mentalizing network. The present
article explores the notion that the understanding of social competitive emotions
is particularly impaired in patients with ventromedial (VM) prefrontal lesions. By
manipulating a simple Theory of Mind (ToM) task, we tested the ability of patients
with localized lesions to understand 'fortune of others' emotions: envy and
gloating (schadenfreude). Patients were also assessed for their ability to
recognize control physical and identification conditions. While envy is an example
of a negative experience in the face of another's fortunes, gloat is thought to be a
positive experience in the face of another's misfortune. Whereas in
schadenfreude and envy the emotion of the self and the protagonist may be
opposite, identification involves matching between the protagonist's and the
observer's emotions. Patients with VM (N = 10) lesions (particularly in the right
hemisphere), although showing intact performance on a basic first order ToM
condition, and relatively preserved understanding of identification, did not
recognize envy (F[6,76] = 3.491, P = 0.004) and gloating (F[6,76] = 3.738, P =
0.003). Impaired recognition of gloating involved additionally lesions in the
inferior parietal lobule (P = 0.001). Furthermore, while patients with lesions in the
left hemisphere were more impaired in recognizing gloating (a positive emotion),
right hemisphere patients were more impaired in recognizing envy (a negative
emotion), suggesting that the valence of these emotions may also be affected by
the asymmetry of the lesion (F[6,68] = 2.002, P = 0.011). In addition, the ability to
identify these emotions was related to perspective-taking abilities and ToM. We
suggest that these results indicate that the mentalizing network including the VM
has a fundamental role in mediating the understanding of competitive emotions
such as envy and gloating.

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Engels AS, Heller W, Mohanty A, Herrington JD, Banich MT, Webb AG,
Miller GA. Specificity of regional brain activity in anxiety types during emotion
processing. Psychophysiology. 2007 May;44(3):352-63

The present study tested the hypothesis that anxious apprehension involves
more left- than right-hemisphere activity and that anxious arousal is associated
with the opposite pattern. Behavioral and fMRI responses to threat stimuli in an
emotional Stroop task were examined in nonpatient groups reporting anxious
apprehension, anxious arousal, or neither. Reaction times were longer for
negative than for neutral words. As predicted, brain activation distinguished
anxious groups in a left inferior frontal region associated with speech production
and in a right-hemisphere inferior temporal area. Addressing a second
hypothesis about left-frontal involvement in emotion, distinct left frontal regions
were associated with anxious apprehension versus processing of positive
information. Results support the proposed distinction between the two types of
anxiety and resolve an inconsistency about the role of left-frontal activation in
emotion and psychopathology.

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Wildgruber D, Ackermann H, Kreifelts B, Ethofer T. Cerebral processing of
linguistic and emotional prosody: fMRI studies. Prog Brain Res. 2006;156:249-
68.

During acoustic communication in humans, information about a speaker's

emotional state is predominantly conveyed by modulation of the tone of voice
(emotional or affective prosody). Based on lesion data, a right hemisphere
superiority for cerebral processing of emotional prosody has been assumed.
However, the available clinical studies do not yet provide a coherent picture with
respect to interhemispheric lateralization effects of prosody recognition and
intrahemispheric localization of the respective brain regions. To further delineate
the cerebral network engaged in the perception of emotional tone, a series of
experiments was carried out based upon functional magnetic resonance imaging
(fMRI). The findings obtained from these investigations allow for the separation of
three successive processing stages during recognition of emotional prosody: (1)
extraction of suprasegmental acoustic information predominantly subserved by
right-sided primary and higher order acoustic regions; (2) representation of
meaningful suprasegmental acoustic sequences within posterior aspects of the
right superior temporal sulcus; (3) explicit evaluation of emotional prosody at the
level of the bilateral inferior frontal cortex. Moreover, implicit processing of
affective intonation seems to be bound to subcortical regions mediating
automatic induction of specific emotional reactions such as activation of the
amygdala in response to fearful stimuli. As concerns lower level processing of
the underlying suprasegmental acoustic cues, linguistic and emotional prosody
seem to share the same right hemisphere neural resources. Explicit judgment of
linguistic aspects of speech prosody, however, appears to be linked to left-sided
language areas whereas bilateral orbitofrontal cortex has been found involved in
explicit evaluation of emotional prosody. These differences in hemispheric
lateralization effects might explain that specific impairments in nonverbal
emotional communication subsequent to focal brain lesions are relatively rare
clinical observations as compared to the more frequent aphasic disorders.

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Sander K, Scheich H Left auditory cortex and amygdala, but right insula
dominance for human laughing and crying. J Cogn Neurosci 2005
Oct;17(10):1519-31.

Evidence suggests that in animals their own species-specific communication

sounds are processed predominantly in the left hemisphere. In contrast,
processing linguistic aspects of human speech involves the left hemisphere,
whereas processing some prosodic aspects of speech as well as other not yet
well-defined attributes of human voices predominantly involves the right
hemisphere. This leaves open the question of hemispheric processing of
universal (species-specific) human vocalizations that are more directly
comparable to animal vocalizations. The present functional magnetic resonance
imaging study addresses this question. Twenty subjects listened to human
laughing and crying presented either in an original or time-reversed version while
performing a pitch-shift detection task to control attention. Time-reversed
presentation of these sounds is a suitable auditory control because it does not
change the overall spectral content. The auditory cortex, amygdala, and insula in
the left hemisphere were more strongly activated by original than by time-
reversed laughing and crying. Thus, similar to speech, these nonspeech
vocalizations involve predominantly left-hemisphere auditory processing.
Functional data suggest that this lateralization effect is more likely based on
acoustical similarities between speech and laughing or crying than on similarities
with respect to communicative functions. Both the original and time-reversed
laughing and crying activated more strongly the right insula, which may be
compatible with its assumed function in emotional self-awareness.

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Noesselt T, Driver J, Heinze HJ, Dolan R. Asymmetrical activation in the

human brain during processing of fearful faces. Carr Biol. 2005 Mar 8;15(5):424-
9.

Traditional split-field studies and patient research indicate a privileged role for the
right hemisphere in emotional processing [1-7], but there has been little direct
fMRI evidence for this, despite many studies on emotional-face processing [8-
10](see Supplemental Background). With fMRI, we addressed differential
hemispheric processing of fearful versus neutral faces by presenting subjects
with faces bilaterally [11-13]and orthogonally manipulating whether each
hemifield showed a fearful or neutral expression prior to presentation of a
checkerboard target. Target discrimination in the left visual field was more
accurate after a fearful face was presented there. Event-related fMRI showed
right-lateralized brain activations for fearful minus neutral left-hemifield faces in
right visual areas, as well as more activity in the right than in the left amygdala.
These activations occurred regardless of the type of right-hemifield face shown
concurrently, concordant with the behavioral effect. No analogous behavioral or
fMRI effects were observed for fearful faces in the right visual field (left
hemisphere). The amygdala showed enhanced functional coupling with right-
middle and anterior-fusiform areas in the context of a left-hemifield fearful face.
These data provide behavioral and fMRI evidence for right-lateralized emotional
processing during bilateral stimulation involving enhanced coupling of the
amygdala and right-hemispheric extrastriate cortex.

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 Wildgruber D Riecker A, Hertrich I, Erb M, Grodd W, Ethofer T,
Ackermann H. Identification of emotional intonation evaluated by fMRI.
Neuroimage. 2005 Feb 15;24(4):1233-41.

During acoustic communication among human beings, emotional information can

be expressed both by the propositional content of verbal utterances and by the
modulation of speech melody (affective prosody). It is well established that
linguistic processing is bound predominantly to the left hemisphere of the brain.
By contrast, the encoding of emotional intonation has been assumed to depend
specifically upon right-sided cerebral structures. However, prior clinical and
functional imaging studies yielded discrepant data with respect to
interhemispheric lateralization and intrahemispheric localization of brain regions
contributing to processing of affective prosody. In order to delineate the cerebral
network engaged in the perception of emotional tone, functional magnetic
resonance imaging (fMRI) was performed during recognition of prosodic
expressions of five different basic emotions (happy, sad, angry, fearful, and
disgusted) and during phonetic monitoring of the same stimuli. As compared to
baseline at rest, both tasks yielded widespread bilateral hemodynamic responses
within frontal, temporal, and parietal areas, the thalamus, and the cerebellum. A
comparison of the respective activation maps, however, revealed comprehension
of affective prosody to be bound to a distinct right-hemisphere pattern of
activation, encompassing posterior superior temporal sulcus (Brodmann Area
[BA] 22), dorsolateral (BA 44/45), and orbitobasal (BA 47) frontal areas.
Activation within left-sided speech areas, in contrast, was observed during the
phonetic task. These findings indicate that partially distinct cerebral networks
subserve processing of phonetic and intonational information during speech
perception.

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Lee GP, Meador KJ, Loring DW, Allison JD, Brown WS, Paul LK, Pillai JJ,
Lavin TB. Neural substrates of emotion as revealed by functional magnetic
resonance imaging. Cogn Behav Neuro. 2004 Mar;17(1):9-17.

OBJECTIVES: To examine the brain circuitry involved in emotional experience

and determine whether the cerebral hemispheres are specialized for positive and
negative emotional experience. BACKGROUND: Recent research has provided
a preliminary sketch of the neurologic underpinnings of emotional processing
involving specialized contributions of limbic and cortical brain regions.
Electrophysiologic, functional imaging, and Wada test data have suggested
positive, approach-related emotions are associated with left cerebral hemisphere
regions, whereas negative, withdrawal-related emotions appear to be more
aligned with right hemisphere mechanisms. METHOD: These emotional-neural
associations were investigated using functional magnetic resonance imaging in
10 healthy controls with 20 positively and 20 negatively valenced pictures from
the International Affective Picture System in a counterbalanced order. Pictures
were viewed within a 1.5 Telsa scanner through computerized video goggles.
RESULTS: Emotional pictures resulted in significantly increased blood flow
bilaterally in the mesial frontal lobe/anterior cingulate gyrus, dorsolateral frontal
lobe, amygdala/anterior temporal regions, and cerebellum. Negative emotional
pictures resulted in greater activation of the right hemisphere, and positive
pictures caused greater activation of the left hemisphere. CONCLUSIONS:
Results are consistent with theories emphasizing the importance of circuitry
linking subcortical structures with mesial temporal, anterior cingulate, and frontal
lobe regions in emotion and with the valence model of emotion that posits
lateralized cerebral specialization for positive and negative emotional experience.

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Baas D, Aleman A, Kahn RS Lateralization of amygdala activation: a systematic

review of functional neuroimaging studies. Brain Res Brain Res Rev. 2004
May;45(2):96-103.

Functional neuroimaging studies of emotion processing consistently report

amygdala activation. Most of these studies observed lateralized amygdala
activity, indicative of a clear hemisphere-specific processing difference between
the left and right amygdalae. Because individual studies use varying paradigms
and are limited by statistical power and sensitivity, it has remained unclear
whether the left or the right amygdala is more consistently involved in emotional
processing. By combining results across 54 fMRI and PET studies in a
metaanalysis, we sought to establish if a common pattern of lateralized amygdala
activation exists. Our findings indicate that across studies, the left amygdala is
more often activated than the right amygdala, suggesting different roles for the
left and right amygdalae in emotional processing. Further analysis showed that
this predominant left amygdala activation is not significantly related to stimulus
type, task instructions, differential habituation rates of the left and right
amygdalae or elaborate processing. The results are discussed

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.
Baas D, Aleman A, Kahn RS. Lateralization of amygdala activation: a
systematic review of functional neuroimaging studies. Brain Res Brain Res Rev.
2004 May;45(2):96-103

Functional neuroimaging studies of emotion processing consistently report

amygdala activation. Most of these studies observed lateralized amygdala
activity, indicative of a clear hemisphere-specific processing difference between
the left and right amygdalae. Because individual studies use varying paradigms
and are limited by statistical power and sensitivity, it has remained unclear
whether the left or the right amygdala is more consistently involved in emotional
processing. By combining results across 54 fMRI and PET studies in a
metaanalysis, we sought to establish if a common pattern of lateralized amygdala
activation exists. Our findings indicate that across studies, the left amygdala is
more often activated than the right amygdala, suggesting different roles for the
left and right amygdalae in emotional processing. Further analysis showed that
this predominant left amygdala activation is not significantly related to stimulus
type, task instructions, differential habituation rates of the left and right
amygdalae or elaborate processing. The results are discussed

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.
Vuilleumier P, Harmony JL, Clarke K, Husain M, Driver J, Dolan RJ.Neural
response to emotional faces with and without awareness: event-related fMRI in a
parietal patient with visual extinction and spatial neglect. Neuropsychologia.
2002;40(12):2156-66

This study examined whether differential neural responses are evoked by

emotional stimuli with and without conscious perception, in a patient with visual
neglect and extinction. Stimuli were briefly shown in either right, left, or both
fields during event-related fMRI. On bilateral trials, either a fearful or neutral left
face appeared with a right house, and it could either be extinguished from
awareness or perceived. Seen faces in left visual field (LVF) activated primary
visual cortex in the damaged right-hemisphere and bilateral fusiform gyri.
Extinguished left faces increased activity in striate and extrastriate cortex,
compared with right houses only. Critically, fearful faces activated the left
amygdala and extrastriate cortex both when seen and when extinguished; as well
as bilateral orbitofrontal and intact right superior parietal areas. Comparison of
perceived versus extinguished faces revealed no difference in amygdala for
fearful faces. Conscious perception increased activity in fusiform, parietal and
prefrontal areas of the left-hemisphere, irrespective of emotional expression;
while a differential emotional response to fearful faces occurring specifically with
awareness was found in bilateral parietal, temporal, and frontal areas. These
results demonstrate that amygdala and orbitofrontal cortex can be activated by
emotional stimuli even without awareness after parietal damage; and that
substantial unconscious residual processing can occur within spared brain areas
well beyond visual cortex, despite neglect and extinction.

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Canli T, Desmond JE, Zhao Z, Glover G, Gabrieli JD Hemispheric
asymmetry for emotional stimuli detected with fMRI. Neuroreport. 1998 Oct
5;9(14):3233-9.

Current brain models of emotion processing hypothesize that positive (or

approach-related) emotions are lateralized towards the left hemisphere, whereas
negative (or withdrawal-related) emotions are lateralized towards the right
hemisphere. Brain imaging studies, however, have so far failed to document
such hemispheric lateralization. In a functional magnetic resonance imaging
(fMRI) study, 14 female subjects viewed alternating blocks of emotionally
valenced positive and negative pictures. When the experience of valence was
equated for arousal, overall brain reactivity was lateralized towards the left
hemisphere for positive pictures and towards the right hemisphere for negative
pictures. This study provides direct support for the valence hypothesis, under
conditions of equivalent arousal, by means of functional brain imaging.

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