Affective Neuroscience - Wikipedia

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From Wikipedia, the free encyclopedia

Affective neuroscience is the study of the neural
mechanisms of emotion. This interdisciplinary
field combines neuroscience with the
psychological study of personality, emotion, and
mood.
[1]
Contents [hide]
1 Brain areas related to emotion
1.1 Main structures of the limbic system
1.2 Other brain structures related to emotion
1.3 Role of the Right Hemisphere in Emotion
1.3.1 The Right Hemisphere Hypothesis
1.3.2 The Valence Hypothesis
2 Relationship to cognitive neuroscience
3 Cognitive neuroscience tasks in affective
neuroscience research
3.1 Emotion Go/No-Go
3.2 Emotional Stroop
3.3 Ekman 60 faces task
3.4 Dot probe (emotion)
3.5 Fear potentiated startle
4 Affective neuroscience and learning
5 Meta-analyses of affective neuroscience
5.1 Locationist approaches
5.2 Psychological constructionist approaches
5.3 Phan et al. 2002
5.4 Murphy et al. 2003
5.5 Barrett et al. 2006
5.6 Kober et al. 2008
5.7 Vytal et al. 2010
5.8 Lindquist et al. 2012
6 See also
7 References
8 Further reading
Brain areas related to emotion [edit]
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http://en.wikipedia.org/wiki/Affective_neuroscience
Emotions are thought to be related to activity in brain areas that direct our attention, motivate
our behavior, and determine the significance of what is going on around us. Pioneering work by
Paul Broca (1878),
[2]
J ames Papez (1937),
[3]
and Paul D. MacLean (1952)
[4]
suggested that
emotion is related to a group of structures in the center of the brain called the limbic system,
which includes the hypothalamus, cingulate cortex, hippocampi, and other structures. Research
has shown that limbic structures are directly related to emotion, but non-limbic structures have
been found to be of greater emotional relevance. The following brain structures are currently
thought to be involved in emotion:
[5]
Main structures of the limbic system [edit]
Amygdala The amygdalae are two small, round structures located anterior to the
hippocampi near the temporal poles. The amygdalae are involved in detecting and learning
what parts of our surroundings are important and have emotional significance. They are
critical for the production of emotion, and may be particularly so for negative emotions,
especially fear.
[6]
Multiple studies have shown amygdala activation when perceiving a
potential threat; various circuits allow the amygdala to use related past memories to better
judge the possible threat.
[7]
Thalamus- The thalamus is involved in relaying sensory and motor signals to the cerebral
cortex,
[8]
especially visual stimuli. The thalamus also plays an important role in regulating
states of sleep and wakefulness.
[9]
Hypothalamus- The hypothalamus is located below the thalamus. It plays a role in emotional
responses by synthesizing and releasing neurotransmitters which can effect mood, reward
and arousal.
[10]
Hippocampus - The hippocampus is a structure of the medial temporal lobes that is mainly
involved in memory. It works to form new memories and also connecting different senses
such as visual input, smell or sound to memories. The hippocampus allows memories to be
stored long term and also retrieves them when necessary. It is this retrieval that is used
within the amygdala to help evaluate current affective stimulus.
[11]
Fornix The fornix is the main output pathway from the hippocampus to the mammillary
bodies. It has been identified as a main region in controlling spatial memory functions,
episodic memory and executive functions.
[12]
Mammillary body - Mammillary bodies are important for recollective memory.
[13]
Olfactory bulb- The olfactory bulbs are the first cranial nerves, located on the ventral side of
the frontal lobe. They are involved in olfaction, the perception of odors.
[14]
Cingulate gyrus- The cingulate gyrus is located above the corpus callosum and is usually
considered to be part of the limbic system. The different parts of the cingulate gyrus have
different functions, and are involved with affect, visceromotor control, response selection,
skeletomotor control, visuospatial processing, and in memory access.
[15]
A part of the
cingulate gyrus is the anterior cingulate cortex, that is thought to play a central role in
attention
[16]
and behaviorally demanding cognitive tasks.
[17]
It may be particularly important
with regard to conscious, subjective emotional awareness. This region of the brain may also
play an important role in the initiation of motivated behavior.
[citation needed]
Other brain structures related to emotion [edit]
Basal ganglia - Basal ganglia are groups of neuclei found on either side of the thalamus.
Basal ganglia play an important role in motivation.
[18]
Orbitofrontal cortex - Is a major structure involved in decision making and the influence by
emotion on that decision.
[19]
Prefrontal cortex The term prefrontal cortex refers to the very front of the brain, behind the
forehead and above the eyes. It appears to play a critical role in the regulation of emotion
and behavior by anticipating the consequences of our actions. The prefrontal cortex may
play an important role in delayed gratification by maintaining emotions over time and
organizing behavior toward specific goals.
[20]
Ventral striatum The ventral striatum is a group of subcortical structures thought to play
an important role in emotion and behavior. One part of the ventral striatum called the
nucleus accumbens is thought to be involved in the experience of goal-directed positive
emotion. Individuals with addictions experience increased activity in this area when they
encounter the object of their addiction.
Insula The insular cortex is thought to play a critical role in the bodily experience of
emotion, as it is connected to other brain structures that regulate the bodys autonomic
functions (heart rate, breathing, digestion, etc.). This region also processes taste
information and is thought to play an important role in experiencing the emotion of disgust.
Cerebellum - Recently, there has been a considerable amount of work that describes the
role of the cerebellum in emotion as well as cognition, and a "Cerebellar Cognitive Affective
Syndrome" has been described.
[21]
Both neuroimaging studies as well as studies following
pathological lesions in the cerebellum (such as a stroke) demonstrate that the cerebellum
has a significant role in emotional regulation. Lesion studies
[22]
have shown that cerebellar
dysfunction can attenuate the experience of positive emotions. While these same studies do
not show an attenuated response to frightening stimuli, the stimuli did not recruit structures
that normally would be activated (such as the amydala). Rather, alternative limbic structures
were activated, such as the ventromedial prefrontal cortex, the anterior cingulate gyrus, and
the insula. This may indicate that evolutionary pressure resulted in the development of the
cerebellum as a redundant fear-mediating circuit to enhance survival. It may also indicate a
regulatory role for the cerebellum in the neural response to rewarding stimuli, such as
money,
[23]
drugs of abuse,
[24]
and orgasm.
[25]
Role of the Right Hemisphere in Emotion [edit]
The Right Hemisphere has been proposed over time as being directly involved in the processing
of emotion. Scientific theory regarding the role of the right hemisphere has developed over time
and resulted in several models of emotional functioning. C.K. Mills was one of the first
researchers to propose a direct link between the right hemisphere and emotional processing,
having observed decreased emotional processing in patients with lesions to the right
hemisphere.
[26][27]
Emotion was originally thought to be processed in the limbic system
structures such as the hypothalamus and amygdala.
[28]
As of the late 1980s to early 1990s
however, neocortical structures were shown to have an involvement in emotion.
[29]
These
findings led to the development of the Right Hemisphere Hypothesis and the Valence
Hypothesis.
The Right Hemisphere Hypothesis [edit]
The Right Hemisphere Hypothesis asserts that the right hemisphere of the neocortical
structures is specialized for the expression and perception of emotion.
[30]
The Right
hemisphere has been linked with mental strategies that are nonverbal, synthetic, integrative,
holistic, and Gestalt which makes it ideal for processing emotion.
[29]
The right hemisphere is
more in touch with subcortical systems of autonomic arousal and attention as demonstrated in
patients that have increased spatial neglect when damage is associated to the right brain as
opposed to the left brain.
[31]
Right hemisphere pathologies have also been linked with
abnormal patterns of autonomic nervous system responses.
[32]
These findings would help
signify the relationship of the subcortical brain regions to the right hemisphere as having a
strong connection.
The Valence Hypothesis [edit]
The Valence Hypothesis acknowledges the right hemispheres role in emotion, but asserts that it
is mainly focused on the processing of negative emotions whereas the left hemisphere
processes positive emotions. The mode of processing of the two hemispheres has been the
discussion of much debate. One version suggests the lack of a specific mode of processes,
stating that the right hemisphere is solely negative emotion and the left brain is solely positive
emotion.
[33]
A second version suggests that there is a complex mode of processing that occurs,
specifically that there is a hemispheric specialization for the expressing and experiencing of
emotion, with the right hemisphere predominating in the experiencing of both positive and
negative emotion.
[34][35]
More recently, the frontal lobe has been the focus of a large amount of
research, stating that the frontal lobes of both hemispheres are involved in the emotional state,
while the right posterior hemisphere, the parietal and temporal lobes, is involved in the
processing of emotion.
[36]
Decreased right parietal lobe activity has been associated with
depression
[37]
and increased right parietal lobe activity with anxiety arousal.
[38]
The increasing
understanding of the role the different hemispheres play has led to increasingly complicated
models, all based some way of the original valence model.
[39]
Relationship to cognitive neuroscience [edit]
In its broadest sense, cognition refers to all mental processes. However, the study of cognition
has historically excluded emotion and focused on non-emotional processes (e.g., memory,
attention, perception, action, problem solving and mental imagery).
[40]
As a result, the study of
the neural basis of non-emotional and emotional processes emerged as two separate fields:
cognitive neuroscience and affective neuroscience. The distinction between non-emotional and
emotional processes is now thought to be largely artificial, as the two types of processes often
involve overlapping neural and mental mechanisms.
[41]
Thus, when cognition is taken at its
broadest definition, affective neuroscience could also be called the cognitive neuroscience of
emotion.
Cognitive neuroscience tasks in affective neuroscience research
[edit]
Emotion Go/No-Go [edit]
The emotion go/no-go task has been frequently used to study behavioral inhibition, particularly
emotional modulation of this inhibition.
[42]
A derivation of the original go/no-go paradigm, this
task involves a combination of affective go cues, where the participant must make a motor
response as quickly as possible, and affective no-go cues, where a response must be
withheld. Because go cues are more common, the task is able to measure ones ability to
inhibit a response under different emotional conditions.
[43]
The task is common in tests of emotion regulation, and is often paired with neuroimaging
measures to localize relevant brain function in both healthy individuals and those with affective
disorders.
[42][44][45]
For example, go/no-go studies converge with other methodology to
implicate areas of the prefrontal cortex during inhibition of emotionally valenced stimuli.
[46]
Emotional Stroop [edit]
The emotional Stroop task, an adaptation to the original Stroop, measures attentional bias to
emotional stimuli.
[47][48]
Participants must name the ink color of presented words while ignoring
the words themselves.
[49]
In general, participants have more difficulty detaching attention from
affectively valenced words, than neutral words.
[50][51]
This interference from valenced words is
measured by the response latency in naming the color of neutral words as compared with
emotional words.
[48]
This task has been often used to test selective attention to threatening and other negatively
valenced stimuli, most often in relation to psychopathology.
[52]
Disorder specific attentional
biases have been found for a variety of mental disorders.
[52][53]
For example, participants with
spider phobia show a bias to spider-related words but not other negatively valenced words.
[54]
Similar findings have been attributed to threat words related to other anxiety disorders.
[52]
However, other studies have questioned these findings. In fact, anxious participants in some
studies show the Stroop interference effect for both negative and positive words, when the
words are matched for emotionality.
[55][56]
This means that the specificity effects for various
disorders may be largely attributable to the semantic relation of the words to the concerns of
the disorder, rather than simply the emotionality of the words.
[52]
Ekman 60 faces task [edit]
The Ekman faces task is used to measure emotion recognition of six basic emotions.
[57][58]
Black and white photographs of 10 actors (6 male, 4 female) are presented, with each actor
displaying each basic emotion. Participants are usually asked to respond quickly with the name
of the displayed emotion. The task is a common tool to study deficits in emotion regulation in
patients with dementia, Parkinsons, and other cognitively degenerative disorders.
[59]
However,
the task has also been used to analyze recognition errors in disorders such as borderline
personality disorder, schizophrenia, and bipolar disorder.
[60][61][62]
Dot probe (emotion) [edit]
The emotional dot-probe paradigm is a task used to assess selective visual attention to and
failure to detach attention from affective stimuli.
[63][64]
The paradigm begins with a fixation
cross at the center of a screen. An emotional stimulus and a neutral stimulus appear side by
side, after which a dot appears behind either the neutral stimulus (incongruent condition) or the
affective stimulus (congruent condition). Participants asked to indicate when they see this dot,
and response latency is measured. Dots that appear on the same side of the screen as the
image the participant was looking at will be identified more quickly. Thus, it is possible to
discern which object the participant was attending to by subtracting the reaction time to
respond to congruent versus incongruent trials.
[63]
The best documented research with the dot probe paradigm involves attention to threat related
stimuli, such as fearful faces, in individuals with anxiety disorders. Anxious individuals tend to
respond more quickly to congruent trials, which may indicate vigilance to threat and/or failure to
detach attention from threatening stimuli.
[63][65]
A specificity effect of attention has also been
noted, with individuals attending selectively to threats related to their particular disorder. For
example, those with social phobia selectively attend to social threats but not physical
threats.
[66]
However, this specificity may be even more nuanced. Participants with obsessive-
compulsive disorder symptoms initially show attentional bias to compulsive threat, but this bias
is attenuated in later trials due to habituation to the threat stimuli.
[67]
Fear potentiated startle [edit]
Fear-potentiated startle (FPS) has been utilized as a psychophysiological index of fear reaction
in both animals and humans.
[68]
FPS is most often assessed through the magnitude of the
eyeblink startle reflex, which can be measured by electromyography.
[69]
This eyeblink reflex is
an automatic defensive reaction to an abrupt elicitor, making it an objective indicator of fear.
[70]
Typical FPS paradigms involve bursts of noise or abrupt flashes of light transmitted while an
individual attends to a set of stimuli.
[70]
Startle reflexes have been shown to be modulated by
emotion. For example, healthy participants tend to show enhanced startle responses while
viewing negatively valenced images and attenuated startle while viewing positively valenced
images, as compared with neutral images.
[71][72]
The startle response to a particular stimulus is greater under conditions of threat.
[73]
A common
example given to indicate this phenomenon is that ones startle response to a flash of light will
be greater when walking in a dangerous neighborhood at night than it would under safer
conditions. In laboratory studies, the threat of receiving shock is enough to potentiate startle,
even without any actual shock.
[74]
Fear potentiated startle paradigms are often used to study fear learning and extinction in
individuals with posttraumatic stress disorder and other anxiety disorders.
[75][76][77]
In fear
conditioning studies, an initially neutral stimulus is repeatedly paired with an aversive one,
borrowing from classical conditioning.
[78]
FPS studies have demonstrated that PTSD patients
have enhanced startle responses during both danger cues and neutral/safety cues as
compared with healthy participants.
[78][79]
Affective neuroscience and learning [edit]
There are many ways affect plays a role during learning. Recently, affective neuroscience has
done much to discover this role. Deep, emotional attachment to a subject area allows a deeper
[80]
understanding of the material and therefore, learning occurs and lasts. When reading, the
emotions one is feeling in comparison to the emotions being portrayed in the content affects
ones comprehension. Someone who is feeling sad will understand a sad passage better than
someone feeling happy.
[81]
Therefore, a students emotion plays a big role during the learning
process.
Emotion can also be embodied or perceived from words read on a page or a persons facial
expression. Neuroimaging studies using fMRI have demonstrated that the same area of the
brain being activated when one is feeling disgust is also activated when one observes another
person feeling disgust.
[82]
In a traditional learning environment, the teacher's facial expression
can play a critical role in students' language acquisition. Showing a fearful facial expression
when reading passages that contain fearful tones facilitates students learning of the meaning of
certain vocabulary words and comprehension of the passage.
[83]
Meta-analyses of affective neuroscience [edit]
A meta-analysis is a statistical approach to synthesizing results across multiple studies. Several
meta-analyses examining the brain basis of emotion have been conducted. In each meta-
analysis, studies were included that investigate healthy, unmedicated adults and that used
subtraction analysis to examine the areas of the brain that were more active during emotional
processing that during a neutral control condition. The meta-analyses to date predominantly
focus on two theoretical approaches, locationist approaches and psychological construction
approaches.
Locationist approaches [edit]
These approaches to emotion hypothesize that several emotion categories (including
happiness, sadness, fear, anger, and disgust) are biologically basic.
[84][85]
In this view,
emotions are inherited biologically based modules that cannot be broken down into more basic
psychological components.
[84][85][86]
Models following a locationist approach to emotion
hypothesize that all mental states belonging to a single emotional category can be consistently
and specifically localized to either a distinct brain region or a defined networks of brain
regions.
[85][87]
Each basic emotion category also shares other universal characteristics: distinct
facial behavior, physiology, subjective experience and accompanying thoughts and
memories.
[84]
Psychological constructionist approaches [edit]
This approach to emotion hypothesizes that emotions like happiness, sadness, fear, anger and
disgust (and many others) are constructed mental states that occur when many different
systems in the brain work together.
[88]
In this view, networks of brain regions underlie
psychological operations (e.g., language, attention, etc.) that interact to produce many different
kinds of emotion, perception, and cognition.
[89]
One psychological operation critical for emotion
is the network of brain regions that underlie valence (feeling pleasant/unpleasant) and arousal
(feeling activated and energized).
[88]
Emotions emerge when neural systems underlying
different psychological operations interact (not just those involved in valence and arousal),
producing distributed patterns of activation across the brain. Because emotions emerge from
more basic components, there is heterogeneity within each emotion category; for example, a
person can experience many different kinds of fear, which feel differently, and which
correspond to different neural patterns in the brain. Thus, this view presents a different
approach to understanding the neural bases of emotion than locationist approaches.
Phan et al. 2002 [edit]
In the first neuroimaging meta-analysis of emotion, Phan et al. (2002) analyzed the results of 55
studies published in peer reviewed journal articles between J anuary 1990 and December 2000
to determine if the emotions of fear, sadness, disgust, anger, and happiness were consistently
associated with activity in specific brain regions. All studies used fMRI or PET techniques to
investigate higher-order mental processing of emotion (studies of low-order sensory or motor
processes were excluded). The authors analysis approach was to tabulate the number of
studies that reported activation in specific brain regions during tasks inducing fear, sadness,
disgust, anger, and happiness. For each brain region, statistical chi-squared analysis was
conducted to determine if the proportion of studies reporting activation during one emotion was
significantly higher than the proportion of studies reporting activation during the other emotions.
Two regions showed this statistically significant pattern across studies. In the amygdala, 66% of
studies inducing fear reported activity in this region, as compared to ~20% of studies inducing
happiness, ~15% of studies inducing sadness (with no reported activations for anger or
disgust). In the subcallosal cingulate, 46% of studies inducing sadness reported activity in this
region, as compared to ~20% inducing happiness and ~20% inducing anger. This pattern of
clear discriminability between emotion categories was in fact rare, with a number of other
patterns occurring in limbic regions (including amydala, hippocampus, hypothalamus, and
orbitofrontal cortex), paralimbic regions (including subcallosal cingulate, medial prefrontal
cortex, anterior cingulate cortex, posterior cingulate cortex, insula, and temporal pole), and
uni/heteromodal regions (including lateral prefrontal cortex, primary sensorimotor cortex,
temporal cortex, cerebellum, and brainstem). Brain regions implicated across discrete emotion
included the basal ganglia (~60% of studies inducing happiness and ~60% of studies inducing
disgust reported activity in this region) and medial prefrontal cortex (happiness ~60%, anger
~55%, sadness ~40%, disgust ~40%, and fear ~30%).
[90]
Murphy et al. 2003 [edit]
Murphy, et al. 2003 analyzed 106 peer reviewed journals published between J anuary 1994 and
December 2001 to examine the evidence for regional specialization of discrete emotions (fear,
disgust, anger, happiness and sadness) across a larger set of studies that Phan et al. Studies
included in the meta-analysis measured activity in the whole brain and regions of interest
(activity in individual regions of particular interest to the study). 3-D Kolmogorov-Smirnov (KS3)
statistics were used to compare rough spatial distributions of 3-D activation patterns to
determine if statistically significant activations (consistently activated across studies) were
specific to particular brain regions for all emotional categories. This pattern of consistently
activated, regionally specific activations was identified in four brain regions: amygdala with fear,
insula with disgust, globus pallidus with disgust, and lateral orbitofrontal cortex with anger. The
amygdala was consistently activated in ~40% of studies inducing fear, as compared to less
than 20% studies inducing happiness, sadness, or anger. The insula was consistently activated
in ~70% of studies inducing disgust, as compared to sadness (~40%), anger (~20%), fear
(~20%), and happiness (~10%). Similar to the insula, the globus pallidus was consistently
activated in ~70% of studies inducing disgust, as compared to less than 25% of studies
inducing sadness, fear, anger or happiness. The lateral orbitofrontal cortex was consistently
activated in over 80% of studies inducing anger, as compared to fear (~30%), sadness (~20%),
happiness (<20%) and disgust (<20%). Other regions showed different patterns of activation
across categories. For example, both the dorsal medial prefrontal cortex and the rostral anterior
cingulate cortex showed consistent activity across emotions (happiness ~50%, sadness ~50%,
anger ~40%, fear ~30%, and disgust ~20%).
[91]
Barrett et al. 2006 [edit]
Barrett, et al. 2006 examined 161 studies published between 1990-2001, subsets of which were
analyzed in previous meta-analyses (Phan, et al. 2002 and Murphy et al. 2003). In this review,
the authors examined the locationist hypothesis by comparing the consistency and specificity of
prior meta-analytic findings specific to each hypothesized basic emotion (fear, anger, sadness,
disgust, and happiness). Consistent neural patterns were defined by brain regions showing
increased activity for a specific emotion (relative to a neutral control condition), regardless of
the method of induction used (for example, visual vs. auditory cue). Specific neural patterns
were defined as architecturally separate circuits for one emotion vs. the other emotions (for
example, the fear circuit must be discriminable from the anger circuit, although both circuits
may include common brain regions). In general, the results supported consistency among the
findings of Phan et al. and Murphy et al., but not specificity. Consistency was determined
through the comparison of chi-squared analyses that revealed whether the proportion of studies
reporting activation during one emotion was significantly higher than the proportion of studies
reporting activation during the other emotions. Specificity was determined through the
comparison of emotion-category brain-localizations by contrasting activations in key regions
that were specific to particular emotions. Increased amygdala activation during fear was the
most consistently reported across induction methods (but not specific). Both meta-analyses
also reported increased activations in regions of the anterior cingulate cortex during sadness,
although this finding was less consistent (across induction methods) and was not specific to
sadness. Both meta-analyses also found that disgust was associated with increased activity in
the basal ganglia, but these findings were neither consistent nor specific. Neither consistent nor
specific activity was observed across the meta-analyses for anger or for happiness. This meta-
analysis additionally introduced the concept of the basic, irreducible elements of emotional life
as dimensions such as approach and avoidance. This dimensional approach involved in
psychological constructionist approaches is further examined in later meta-analyses of Kober et
al. 2008 and Lindquist et al. 2012.
[88]
Kober et al. 2008 [edit]
Instead of investigating specific emotions, Kober, et al. 2008 reviewed 162 neuroimaging
studies published between 1990-2005 to determine if groups of brain regions show consistent
patterns of activation during emotional experience (that is, actively experiencing an emotion
first-hand) and during emotion perception (that is, perceiving a given emotion as experienced
by another). This meta-analysis used multilevel kernal density analysis (MKDA) to examine
fMRI and PET studies, a technique that prevents single studies from dominating the results
(particularly if they report multiple nearby peaks) and that enables studies with large sample
sizes (those involving more participants) to exert more influence upon the results. MKDA was
used to establish a neural reference space that includes the set of regions showing consistent
increases across all studies (for further discussion of MDKA see Wager et al. 2007).
[92]
Next,
this neural reference space was partitioned into functional groups of brain regions showing
similar activation patterns across studies by first using multivariate techniques to determine co-
activation patterns and then using data-reduction techniques to define the functional groupings
(resulting in six groups). Consistent with a psychological construction approach to emotion, the
authors discuss each functional group in terms more basic psychological operations. The first
Core Limbic group included the left amygdala, hypothalamus, periaqueductal gray/thalamus
regions, and amygdala/ventral striatum/ventral globus pallidus/thalamus regions, which the
authors discuss as an integrative emotional center that plays a general role in evaluating
affective significance. The second Lateral Paralimbic group included the ventral anterior
insula/frontal operculum/right temporal pole/ posterior orbitofrontal cortex, the anterior insula/
posterior orbitofrontal cortex, the ventral anterior insula/ temporal cortex/ orbitofrontal cortex
junction, the midinsula/ dorsal putamen, and the ventral striatum /mid insula/ left hippocampus,
which the authors suggest plays a role in motivation, contributing to the general valuation of
stimuli and particularly in reward. The third Medial Prefrontal Cortex group included the dorsal
medial prefrontal cortex, pregenual anterior cingulate cortex, and rostral dorsal anterior
cingulate cortex, which the authors discuss as playing a role in both the generation and
regulation of emotion. The fourth Cognitive/ Motor Network group included right frontal
operculum, the right interior frontal gyrus, and the pre-supplementray motor area/ left interior
frontal gyrus, regions that are not specific to emotion, but instead appear to play a more
general role in information processing and cognitive control. The fifth Occipital/ Visual
Association group included areas V8 and V4 of the primary visual cortex, the medial temporal
lobe, and the lateral occipital cortex, and the sixth Medial Posterior group included posterior
cingulate cortex and area V1 of the primary visual cortex. The authors suggest that these
regions play a joint role in visual processing and attention to emotional stimuli.
[93]
Vytal et al. 2010 [edit]
Vytal, et al. 2010 examined 83 neuroimaging studies published between 1993-2008 to examine
whether neuroimaging evidence supports the idea of biologically discrete, basic emotions (i.e.
fear, anger, disgust, happiness, and sadness). Consistency analyses identified brain regions
that were associated with a given emotion. Discriminability analyses identified brain regions
that were significantly, differentially active when contrasting pairs of discrete emotions. This
meta-analysis examined PET or fMRI studies that reported whole brain analyses identifying
significant activations for at least one of the five emotions relative to a neutral or control
condition. The authors used activation likelihood estimation (ALE) to perform spatially sensitive,
voxel-wise (sensitive to the spatial properties of voxels) statistical comparisons across studies.
This technique allows for direct statistical comparison between activation maps associated with
each discrete emotion. Thus, discriminability between the five discrete emotion categories was
assessed on a more precise spatial scale than what had been accomplished in prior meta-
analyses. Consistency was first assessed by comparing the ALE map generated across studies
for each emotion (for example, the ALE map identifying regions consistently activated by
studies inducing fear) to ALE map generated by random permutations. Discriminability was
then assessed by pair-wise contrasts of individual emotion ALE maps (for example, fear ALE
map vs. anger ALE map; fear ALE map vs. disgust map) across all basic emotions pairings.
Consistent and discriminable patterns of neural activation were observed for the five emotional
categories. Happiness was consistently associated with activity in 9 regional brain clusters, the
largest located in the right superior temporal gyrus. For the first time, happiness was
discriminated from the other emotional categories, with the largest clusters of activity specific to
happiness (vs. the other emotion categories) located in right superior temporal gyrus and left
rostral anterior cingulate cortex. Sadness was consistently associated with 35 clusters (the
largest activation cluster located in the left medial frontal gyrus) and was discriminated from the
other emotion categories by significantly greater activity in left medial frontal gyrus, right middle
temporal gyrus, and right inferior frontal gyrus. Anger was consistently associated with activity
in 13 clusters (the largest of which was located in the left inferior frontal gyrus), and was
discriminated from the other emotion categories by significantly greater activity in bilateral
inferior frontal gyrus, and in right parahippocampal gyrus. Fear was consistently associated
with 11 clusters (the largest activation cluster in the left amygdala) and was discriminated from
the other emotion categories by significantly greater activity in the left amygdala and left
putamen. Disgust was consistently activated with 16 clusters (the largest activation cluster in
the right insula/ right inferior frontal gyrus) and was discriminated from the other emotion
categories by significantly greater activity in the right putamen and the left insula.
[94]
Lindquist et al. 2012 [edit]
Lindquist, et al. 2012 reviewed 91 PET and fMRI studies published between J anuary 1990 and
December 2007. The studies included in this meta-analysis used induction methods that elicit
emotion experience or emotion perception of fear, sadness, disgust, anger, and happiness. The
goal was to compare locationist approaches with psychological constructionist approaches to
emotion. Similar to Kober et al. described above, a Multilevel Peak Kernel Density Analysis
[92]
transformed the individual peak activations reported across study contrasts into a neural
reference space (in other words, the set of brain regions consistently active across all study
contrasts assessing emotion experience or perception). The density analysis was then used to
identify regions (or voxels) within the neural reference space with more consistent activations
for a specific emotion category (anger, fear, happiness, sadness, and disgust) than all other
emotions. Chi-squared analysis was used to create statistical maps that indicated if each
previously identified and consistently active regions (those identified during density analysis)
were more frequently activated in studies of each emotion category versus the average of all
other emotions, regardless of activations elsewhere in the brain. Chi-squared analysis and
density analysis both defined functionally consistent and selective regions, or regions which
showed a relatively more consistent increase in activity for the experience or perception of one
emotion category across studies in the literature. Thus, a selective region could present
increased activations relatively more so to one emotion category while also having a response
to multiple other emotional categories. A series of logistic regressions were then preformed to
identify if any of the regions that were identified as consistent and selective to an emotion
category were additionally specific to a given category. Regions were defined as specific to a
given emotion if they showed increased activations for only one emotional category, and never
showed increased activity during instances of the other emotional categories. In other words, a
region could be defined as consistent, selective and specific for e.g. fear perception if it only
showed significantly greater increases in activation during the perception of fear and did not
show increased activity during any other emotion categories. However, the same region would
be defined as only consistent and selective (and not specific) to fear perception if it additionally
displayed increased activations during anger perception. Strong support for the locationist
approach was defined as evidence that basic emotion categories (anger, disgust, fear,
happiness and sadness) consistently map onto areas of the brain that specifically activate in
response to instances of only one emotional category. Strong support for the constructionist
approach was defined as evidence that multiple psychological operations (some of which are
not specific or selective to emotion) consistently occur across many brain regions and multiple
emotional categories.
The results indicated that many brain regions demonstrated consistent and selective activations
in the experience or perception of an emotion category (versus all the other emotion
categories). Consistent with constructionist models, however, no region demonstrated
functional specificity for the emotions of fear, disgust, happiness, sadness or anger. Based on
the existing scientific literature, the authors proposed different roles for the brain regions that
have traditionally been associated with only one emotion category. The authors propose that
the amygdala, anterior insula, orbitofrontal cortex each contribute to core affect, which are
basic feelings that are pleasant or unpleasant with some level of arousal. The amygdala, for
example, appears to play a more general role in indicating if external sensory information is
motivationally salient, and is particularly active when a stimulus is novel or evokes uncertainty.
The anterior insula may represent core affective feelings in awareness across a number of
emotion categories, driven largely by sensations originating in the body. The orbitofrontal cortex
appears to function as a site for integrating sensory information from the body and sensory
information from the world to guide behavior. Closely related to core affect, the authors propose
that anterior cingulate and dorsolateral prefrontal cortex play vital roles in attention, with
anterior cingulate supporting the use of sensory information for directing attention and motor
responses during response selection and with dorsolateral prefrontal cortex supporting
executive attention. In many psychological construction approaches, emotions also involve the
act of interpreting ones situation in the world relative to the internal state of the body, or what is
referred to as conceptualization. In support of this idea, the dorsomedial prefrontal cortex and
hippocampus were consistently active in this meta-analysis, regions that appear to play an
important role conceptualizing during emotion, which are also involved in simulating previous
experience (e.g. knowledge, memory). Language is also central to conceptualizing, and regions
that support language, including ventrolateral prefrontal cortex, were also consistently active
across studies of emotion experience and perception.
[89]
See also [edit]
Affect (psychology)
Affective science
Affective spectrum
Emotion
Endocrinology
List of topics related to brain mapping
Music therapy
Neuroendocrinology
Psychophysiology
Psychology
Psychiatry
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Further reading [edit]
Davidson, R.J ., & Irwin, W. (1999). The functional neuroanatomy of emotion and affective
style. Trends in Cognitive Science, 3, 11-21.
Freitas-Magalhaes, A. (2009). Emotional Expression: The Brain and The Face. Porto:
University Fernando Pessoa Press. ISBN 978-989-643-034-4
Panksepp, J . (1992). A critical role for affective neuroscience in resolving what is basic
about basic emotions" Psychological Review 99, 554-60.
Harmon-J ones E, & Winkielman P. (Eds.) Social Neuroscience: Integrating Biological and
Psychological Explanations of Social Behavior. New York: Guilford Publications.
Cacioppo, J .T., & Berntson, G.G. (2005). Social Neuroscience. Psychology Press.
Cacioppo, J .T., Tassinary, L.G., & Berntson, G.G. (2007). Handbook of Psychophysiology.
Cambridge University Press.
Panksepp J . (1998). Affective Neuroscience: The Foundations of Human and Animal
Emotions (Series in Affective Science). Oxford University Press, New York, New York.
Brain and Cognition, Vol. 52, No. 1, pp. 1133 (J une, 2003). Special Issue on Affective
Neuroscience.
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