Brain and Cognition xxx (2015) xxx–xxx

Contents lists available at ScienceDirect

Brain and Cognition

journal homepage: www.elsevier.com/locate/b&c

Affective mapping: An activation likelihood estimation (ALE)

meta-analysis
Lauren A.J. Kirby a,⇑, Jennifer L. Robinson a,b,c
a
Department of Psychology, Auburn University, Auburn, AL 36849, United States
b
Department of Electrical and Computer Engineering, Auburn University, Auburn, AL 36849, United States
c
Department of Kinesiology, Auburn University, Auburn, AL 36849, United States

a r t i c l e i n f o a b s t r a c t

Article history: Functional neuroimaging has the spatial resolution to explain the neural basis of emotions. Activation
Accepted 24 April 2015 likelihood estimation (ALE), as opposed to traditional qualitative meta-analysis, quantifies convergence
Available online xxxx of activation across studies within affective categories. Others have used ALE to investigate a broad range
of emotions, but without the convenience of the BrainMap database. We used the BrainMap database and
Keywords: analysis resources to run separate meta-analyses on coordinates reported for anger, anxiety, disgust, fear,
Neuroimaging happiness, humor, and sadness. Resultant ALE maps were compared to determine areas of convergence
Emotion
between emotions, as well as to identify affect-specific networks. Five out of the seven emotions
Activation likelihood estimation
Brainmap
demonstrated consistent activation within the amygdala, whereas all emotions consistently activated
Affect programs the right inferior frontal gyrus, which has been implicated as an integration hub for affective and
fMRI cognitive processes. These data provide the framework for models of affect-specific networks, as well
as emotional processing hubs, which can be used for future studies of functional or effective connectivity.
Ó 2015 Elsevier Inc. All rights reserved.

1. Introduction of emotion see Lindquist, Wager, Kober, Bliss-Moreau, & Barrett,

Emotions are biologically based motivating states causing
changes in feeling, behavior, and neurological and physiological
function (Barrett, 2012; Barrett, Mesquita, Ochsner, & Gross,
2007) and likely have adaptive functions for communicating and
making decisions (Ekman, 1999; Nesse & Ellsworth, 2009; Shariff
& Tracy, 2011). Understanding brain function involved in affective
processing could elucidate the neurophysiological mechanisms
associated with the experience of emotions and potentially gener-
ate a biosignature of discrete affective states for reference in a vari-
ety of basic and applied research contexts. Neuroimaging provides
the best method in an array of converging evidence to better
understand the internal mechanisms associated with emotions.
Many researchers have undertaken the task of mapping the neural
activity associated with the experience of various emotions (for an
early qualitative meta-analysis see Davidson & Irwin, 1999; for
quantitative meta-analyses, see Fusar-Poli et al., 2009; Kober
et al., 2008; Murphy, Nimmo-Smith, & Lawrence, 2003; Phan,
Wager, Taylor, & Liberzon, 2002; and Wager, Phan, Liberzon, &
Taylor, 2003, and for a comprehensive review of the neural basis
⇑ Corresponding author at: Department of Psychology, Auburn University, 226
Thach Hall, Auburn, AL 36849, United States.
E-mail addresses: lajkirby@auburn.edu (L.A.J. Kirby), jrobinson@auburn.edu
(J.L. Robinson).
2012). However, there is variation in the models of emotion exam-
ined as well as the meta-analytic techniques used. Activation like-
lihood estimation (ALE) is a meta-analytic method that is arguably
best suited to address this issue, as discussed below.
1.1. Previous reviews
Previous meta-analyses on the neuroimaging of emotion have
examined varied models of emotions. Murphy et al. (2003)
describe and test single-system, dual-system (dimensional, such
as valence or motivation), and multi-system models of emotion.
To test across single system models, they used the 3-D
Kolmogorov–Smirnov statistic (KS3; a non-parametric test
between two distributions) across emotion conditions and hemi-
spheric differences. When testing a multi-system model, regional
specialization for discrete emotions, they found the most consis-
tently activated region for each emotion investigated (fear, disgust,
anger, happiness, and sadness), defined as the brain region
reported active in the largest portion of studies for each discrete
emotion. The anterior cingulate cortex (ACC) and amygdala
showed activation common across emotions. Minimal support for
the limbic system hypothesis (Maclean, 1949) was also found.
Dual-system models failed to find any substantial support. Most
notably a multi-system model of affect programs (basic emotions;

http://dx.doi.org/10.1016/j.bandc.2015.04.006
0278-2626/Ó 2015 Elsevier Inc. All rights reserved.

Please cite this article in press as: Kirby, L. A. J., & Robinson, J. L. Affective mapping: An activation likelihood estimation (ALE) meta-analysis. Brain and
Cognition (2015), http://dx.doi.org/10.1016/j.bandc.2015.04.006
2 L.A.J. Kirby, J.L. Robinson / Brain and Cognition xxx (2015) xxx–xxx

Ekman, 1999) was supported by their strong findings of fear being activation likelihood estimation (ALE) as opposed to traditional
specialized to the amygdala, disgust to the insula and globus pal- qualitative meta-analyses, offer the opportunity to quantify conver-
lidus, and anger to the lateral orbitofrontal cortex (OFC). gence of activation across studies, within affective categories.
More recently, Lindquist et al. (2012) framed their investigation Furthermore, advancements in the coding, storage, search, and
not in terms of particular models of emotion, but as an approach to availability of neuroimaging metadata has been afforded by the
neuroscience in general. They pit locationism, the belief that psy- BrainMap database (Laird, Lancaster, & Fox, 2005). To date, very
chological processes can be localized to a specific region, against few studies have compared the underlying neural networks associ-
psychological constructionism, the belief that psychological phe- ated with a diverse range of emotions. These factors converge to
nomena such as emotion are built from more basic processes. make a BrainMap ALE meta-analysis of neuroimaging metadata of
Locationism was operationally defined as activation within an emotion activation both convenient and pertinent to carry out.
emotion category consistently and uniquely found in a certain Here, we examine the seven emotional categories in the BrainMap
region or set of regions. Constructionism was operationalized as database to determine primary emotional network nodes, which
brain regions being involved in several emotion categories and may provide a more accurate framework for future affective neuro-
grounded in a more basic psychological process. The researchers science studies.
used regions identified in previous density analyses to test the
locationist view, including the amygdala, the anterior insula, the
2. Methods
OFC, the ACC, the dorsomedial prefrontal cortex (DMPFC), medial
temporal lobe (MTL), posterior cingulate cortex (PCC), anterior
2.1. Materials
temporal lobe, ventrolateral PFC, dorsolateral PFC, periaqueductal
gray, the and the visual cortex. They found lack of consistency
The BrainMap database involves three programs: Scribe, Sleuth,
and specificity within emotion categories across studies in the
and GingerALE (Eickhoff et al., 2009; Laird et al., 2005). Scribe is used
queried areas and so concluded the locationist approach (affect
to code neuroimaging studies and upload them to the database.
programs) to be overly reductionistic, favoring the psychological
Sleuth is used for searching the database with a variety of filters
constructionist view of emotions.
(in the form of drop-down menus for options like context, subjects,
While Lindquist and colleagues failed to find consistency and
behavioral domain, and paradigm class) and with features to save
specificity associated with discrete emotions categories, Vytal
and export the results. It currently features the (x, y, z) coordinate
and Hamann (2010) used activation likelihood estimation (ALE)
results of 2396 neuroimaging papers, 105 paradigm classes,
and found consistent activation for happiness, sadness, anger, fear,
46,479 subjects, 11,397 experiments and 91,251 locations (www.
and disgust, as well as confirmed their specificity by comparing
brainmap.org). GingerALE uses the metadata exported into text files
emotion categories against each other. However, their study is
by Sleuth to complete ALE analyses, which put out tables of 3-d coor-
markedly limited in including only 30 neuroimaging papers for
dinates of common activations across studies. All Brainmap Sleuth
analysis. Use of a rigorously coded neuroimaging study database
workspaces can be found at http://aucanlab.com/research/data.
could improve the quantity, quality, and representativeness of
the literature used in such a study.
2.2. Procedure
1.2. Meta-analytic technique
Using the drop-down menu options in BrainMap’s Sleuth search
program (Laird et al., 2005) we selected papers for our
In addition to the variety in models of emotion examined, pre-
meta-analysis by filtering our search by (1) normal mapping (as
vious reviews used different meta-analytic techniques as well.
opposed to studies examining effects of gender, aging, disease,
Activation likelihood estimation (ALE) has several advantages over
handedness, drug effects, etc.), (2) activations only (excluding
traditional qualitative neuroimaging meta-analytic techniques.
coordinates with reported deactivations), (3) healthy, normal par-
ALE uses coordinates or foci reported in standard space by individ-
ticipants, and (4) papers coded as emotion as defined in the
ual studies as its input into a meta-analysis, avoiding the arbitrari-
BrainMap lexicon (http://brainmap.org/scribe/BrainMapLex.xls).
ness of qualitative methods. Also, the coordinates can be converted
We queried the BrainMap database for each of the seven emotion
across different standard spaces, making comparison across differ-
classes available, including anger, anxiety, disgust, fear, happiness,
ent standard spaces and neuro-atlases possible. The foci are then
humor, and sadness and separate ALE meta-analyses were run for
weighted by the n of each study. Probability of activation is calcu-
sets of (x, y, z) coordinates. We generated and compared ALE maps
lated based on both individual voxel statistical significance and a
using BrainMap’s GingerALE 2.3 program (Eickhoff et al., 2009) to
minimum cluster size threshold. ALE then reports only the com-
identify areas of convergence in activation across multiple emo-
mon activation across all studies in order to remove any
tions as well as affect-specific areas. Activations were arbitrarily
task-dependency from the results.
determined to be in the same area if all (x, y, z) coordinates were
within 6 mm of each other.
1.3. Rationale

Affective neuroscience is an important area of study for several 3. Results

reasons. For example, it may give insight into our cognitive and
affective evolution and has implications for applied and clinical
work in emotions. It is now thought that emotions have adaptive
functions, including function as a motivating drive state, for
decision-making, and for communicating (Ekman, 1999; Nesse &
Ellsworth, 2009; Shariff & Tracy, 2011). Investigating the neural cor-
relates of emotions can give us new insight into previously opaque
internal emotional processes. Functional neuroimaging, with its
superior spatial resolution, is best suited over other (imaging and
non-imaging) techniques to address questions about the neuro-
science of emotions. Advancements in analysis techniques, such as
3.1. Affect-specific networks
3.1.1. Anger
The anger search yielded 27 papers, 550 subjects, 64 experi-
ments, 90 conditions, 417 and locations. Cortical activation unique
to anger was found in the left and right IFG (BA47), the left lingual
gyrus (BA 18), and the bilateral middle temporal gyri (MTG; BA
21/22) and right STG (BA 41). Subcortical activation included the
bilateral substantia nigra (SN), the left amygdala (BA 47), and the
left insula (BA 46/47) (Tables 2 and 3).

Please cite this article in press as: Kirby, L. A. J., & Robinson, J. L. Affective mapping: An activation likelihood estimation (ALE) meta-analysis. Brain and
Cognition (2015), http://dx.doi.org/10.1016/j.bandc.2015.04.006
 L.A.J. Kirby, J.L. Robinson / Brain and Cognition xxx (2015) xxx–xxx 3

Table 1
Convergent emotion networks organized by lobe and region.

Areas of convergent emotion activation

Lobe Region Emotion BA x y z
Anterior Left Culmen Humor ⁄ 38 38 26
Left Culmen Happiness ⁄ 36 44 20
Frontal Left Inferior Frontal Gyrus-1 Disgust 47 40 22 14
Left Inferior Frontal Gyrus-1,2 Anger ⁄ 34 24 2
Left Inferior Frontal Gyrus-2 Fear 47 46 14 0
Right Inferior Frontal Gyrus-1 Anxiety 9 38 6 26
Right Inferior Frontal Gyrus-1,2 Disgust 9 44 12 22
Right Inferior Frontal Gyrus-2 Fear 9 46 12 28
Right Inferior Frontal Gyrus-3 Happiness 44 54 6 12
Right Inferior Frontal Gyrus-3 Anger 44 52 4 16
Right Inferior Frontal Gyrus-4 Sadness 47 38 20 10
Right Inferior Frontal Gyrus-4 Disgust 47 40 20 10
Right Inferior Frontal Gyrus-4 Anger ⁄ 40 18 12
Right Inferior Frontal Gyrus-5 Anger ⁄ 44 20 2
Right Inferior Frontal Gyrus-5,6 Happiness 47 46 24 6
Right Inferior Frontal Gyrus-6 Humor ⁄ 46 28 8
Left Middle Frontal Gyrus Sadness ⁄ 42 12 28
Left Middle Frontal Gyrus Fear 46 44 18 22
Left Middle Frontal Gyrus Anger 46 46 18 26
Limbic Left Anterior Cingulate Fear ⁄ 12 38 0
Left Anterior Cingulate Anxiety ⁄ 12 40 2
Right Anterior Cingulate Happiness 32 2 48 2
Right Anterior Cingulate Fear 32 4 46 4
Left Cingulate Gyrus Anxiety 32 6 18 28
Left Cingulate Gyrus Fear 32 2 22 28
Left Parahippocampal Gyrus Happiness Amygdala 22 6 12
Left Parahippocampal Gyrus Fear Amygdala 20 4 10
Left Parahippocampal Gyrus Humor Amygdala 18 4 14
Left Parahippocampal Gyrus Anxiety ⁄ 18 2 10
Right Parahippocampal Gyrus-1 Sadness 28 14 4 10
Right Parahippocampal Gyrus-1 Disgust 28 16 4 10
Right Parahippocampal Gyrus-1,2 Fear Amygdala 20 4 12
Right Parahippocampal Gyrus-2 Anger Amygdala 22 4 12
Right Parahippocampal Gyrus-2 Happiness Amygdala 24 2 12
Right Parahippocampal Gyrus-2 Sadness Amygdala 26 4 16
Midbrain Right Mammillary Body Fear ⁄ 2 14 6
Left Red Nucleus Anxiety ⁄ 0 14 4
Occipital Right Inferior Temporal Gyrus Fear ⁄ 46 70 0
Right Inferior Temporal Gyrus Disgust ⁄ 46 72 0
Posterior Right Declive Happiness ⁄ 22 60 12
Right Declive Happiness ⁄ 28 56 18
Left Tuber Humor ⁄ 46 64 24
Left Tuber Happiness ⁄ 44 64 26
Sub-lobar Left Caudate Anxiety Caudate Head 8 4 0
Left Caudate Sadness Caudate Head 2 6 2
Left Claustrum Anxiety ⁄ 34 10 2
Left Claustrum Disgust ⁄ 36 6 8
Left Insula Disgust 13 36 22 0
Left Insula Anxiety 13 34 20 0
Left Insula Fear 13 30 22 0
Right Insula Disgust 13 34 20 2
Right Insula Anxiety 13 36 16 4
Right Insula Fear ⁄ 38 16 2
Left Thalamus Disgust Lateral Posterior Nucleus 14 18 12
Left Thalamus Fear Medial Dorsal Nucleus 8 16 10
Right Thalamus Anxiety Pulvinar 2 28 0
Right Thalamus Fear Pulvinar 2 30 0
Right Thalamus Fear Medial Dorsal Nucleus 4 18 10
Right Thalamus Anxiety Medial Dorsal Nucleus 6 18 6
Temporal Left Fusiform Gyrus-1 Happiness 37 44 50 16
Left Fusiform Gyrus-1 Fear 37 38 46 18
Left Fusiform Gyrus 1, 2 Fear 37 40 54 16
Left Fusiform Gyrus-2 Anger 37 38 56 12
Left Fusiform Gyrus-2 Disgust 37 42 60 10
Right Fusiform Gyrus Anger 37 38 52 16
Right Fusiform Gyrus Disgust 37 42 58 12
Left Superior Temporal Gyrus Happiness 38 48 12 18
Left Superior Temporal Gyrus Anger 38 48 14 16

Please cite this article in press as: Kirby, L. A. J., & Robinson, J. L. Affective mapping: An activation likelihood estimation (ALE) meta-analysis. Brain and
Cognition (2015), http://dx.doi.org/10.1016/j.bandc.2015.04.006
4 L.A.J. Kirby, J.L. Robinson / Brain and Cognition xxx (2015) xxx–xxx

Table 2
Affect-specific networks organized by lobe and region.

Areas of unique activation

Lobe Region Emotion BA x y z
Anterior Left Culmen Disgust ⁄ 26 48 14
Right Culmen Fear ⁄ 38 50 18
Right Culmen Humor ⁄ 26 56 20
Right Culmen Humor ⁄ 26 42 16
Cerebellum⁄ Right Cerebellum⁄ Humor ⁄ 4 76 32
Frontal Left Inferior Frontal Gyrus Happiness 47 30 20 8
Left Inferior Frontal Gyrus Anger 47 46 22 4
Right Inferior Frontal Gyrus Sadness 45 46 24 16
Right Inferior Frontal Gyrus Anger 47 34 32 2
Right Inferior Frontal Gyrus Happiness 47 42 16 10
Left Middle Frontal Gyrus Sadness 9 36 22 32
Left Middle Frontal Gyrus Sadness 47 40 34 6
Right Middle Frontal Gyrus Sadness 9 44 8 38
Right Middle Frontal Gyrus Happiness 10 38 38 16
Right Middle Frontal Gyrus Disgust 46 46 20 22
Left Medial Frontal Gyrus Sadness 9 4 44 28
Left Medial Frontal Gyrus Sadness 9 6 28 28
Left Superior Frontal Gyrus Sadness 9 8 52 24
Right Cingulate Gyrus Disgust 32 4 18 34
Right Precentral Gyrus Sadness 44 50 14 6
Limbic Left Anterior Cingulate Happiness 32 0 40 8
Left Anterior Cingulate Fear 24 2 26 4
Left Anterior Cingulate Sadness 32 8 18 10
Left Anterior Cingulate Fear 32 8 42 10
Right Anterior Cingulate Anxiety 32 14 42 6
Left Cingulate Gyrus Anxiety 31 10 28 38
Right Cingulate Gyrus Fear 32 2 8 40
Right Cingulate Gyrus Anxiety 31 14 34 38
Left Anterior Cingulate Happiness Hippocampus 32 24 14
Left Anterior Cingulate Fear 28 18 24 8
Right Parahippocampal Gyrus Happiness 35 16 28 10
Right Parahippocampal Gyrus Happiness 35 20 20 8
Right Parahippocampal Gyrus Fear Hippocampus 36 16 14
Midbrain Left Red Nucleus Fear ⁄ 2 26 14
Left Substantia Nigra Anger ⁄ 10 24 6
Right Substantia Nigra Anger ⁄ 14 20 4
Occipital Left Fusiform Gyrus Fear 18 22 88 12
Right Fusiform Gyrus Fear 19 30 82 14
Right Fusiform Gyrus Happiness 19 40 74 10
Left Lingual Gyrus Anger 18 20 56 2
Parietal Left Postcentral Gyrus Disgust 3 40 22 48
Left Postcentral Gyrus Anxiety 40 52 28 20
Right Postcentral Gyrus Happiness 40 58 28 22
Right Superior Parietal Lobule Anxiety 7 30 50 62
Posterior Left Declive Fear ⁄ 32 68 16
Left Declive Happiness ⁄ 38 56 16
Right Declive Fear ⁄ 32 64 20
Right Declive Happiness ⁄ 32 80 16
Left Uvula Humor ⁄ 30 72 24
Sub-lobar Left Amygdala Anger 47 34 24 2
Left Claustrum Disgust ⁄ 36 0 8
Left Insula Anger 46 46 18 26
Left Insula Anger 47 46 22 4
Right Insula Anxiety 13 30 16 6
Right Insula Sadness 13 48 12 2
Left Lentiform Nucleus Sadness Medial Globus Pallidus 16 2 8
Left Lentiform Nucleus Humor Putamen 16 6 2
Right Lentiform Nucleus Sadness Putamen 18 4 8
Right Lentiform Nucleus Disgust Putamen 24 4 2
Left Thalamus Disgust ⁄ 6 8 2
Left Thalamus Disgust Medial Dorsal Nucleus 2 18 4
Left Thalamus Sadness ⁄ 0 4 8
Right Thalamus Disgust Mammillary Body 10 16 0
Temporal Left Middle Temporal Gyrus Anger 22 50 42 4
Left Middle Temporal Gyrus Happiness 39 42 70 16
Left Middle Temporal Gyrus Happiness 21 52 4 16
Left Middle Temporal Gyrus Happiness 21 52 4 14
Right Middle Temporal Gyrus Anger 21 50 42 10
Left Superior Temporal Gyrus Fear 42 54 30 18

Please cite this article in press as: Kirby, L. A. J., & Robinson, J. L. Affective mapping: An activation likelihood estimation (ALE) meta-analysis. Brain and
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 L.A.J. Kirby, J.L. Robinson / Brain and Cognition xxx (2015) xxx–xxx 5

Table 2 (continued)

Areas of unique activation

Lobe Region Emotion BA x y z
Left Superior Temporal Gyrus Anxiety 38 44 8 8
Right Superior Temporal Gyrus Anger 41 52 18 6

3.1.2. Anxiety 3.1.3. Disgust

The anxiety search yielded 25 papers, 574 subjects, 59 experi-
ments, 70 conditions, and 453 locations. Cortical activation unique
to anxiety was found in the right CG (BA31), the right superior
parietal lobule (RSPL; BA 7), and the LSTG (BA 38). Subcortical acti-
vation was found in the right insula (BA 13), the RACC (BA32) and
right CG (BA31) (Tables 2 and 4).
The disgust search yielded 41 papers, 735 subjects, 115 experi-
ments, 131 condition, and 798 locations. Cortical activation unique
to disgust was found in the RCG (BA 32), RMFG (BA 46) and the left
postcentral gyrus (LPosG; BA3). Subcortical activation was found in
the right putamen, the right mammillary body, and the left MDN of
the thalamus. Cerebellar activation was found in the left culmen
(Table 5).

Table 3
3.1.4. Fear
Anger-related ALE results organized by Brodmann area.
The fear search yielded 70 papers, 1426 subjects, 194 experi-
Anger ments, 211 conditions, and 1107 locations. Cortical activation
Lobe Region BA x y z unique to fear was found in left and right FG (BA 18/19), and the
Frontal Left Middle Frontal Gyrus 46 46 18 26 LSTG (BA 42). Subcortical activation included the LACC (BA
Left Inferior Frontal Gyrus-1,2 47 34 24 2 24/32), the RCG (BA 32), and the right hippocampus. Cerebellar
Left Inferior Frontal Gyrus 47 46 22 4 activation was found in the right culmen (Table 6).
Right Inferior Frontal Gyrus-3 44 52 4 16
Right Inferior Frontal Gyrus-4 47 40 18 12
Right Inferior Frontal Gyrus-5 47 44 20 2 3.1.5. Happiness
The happiness search yielded 71 papers, 1357 subjects, 161
Limbic Right Parahippocampal Gyrus- Amygdala 22 4 12
2 experiments, 222 condition, and 966 locations. Cortical activation
unique to happiness was found in the bilateral IFG (BA 47),
Midbrain Left Substantia Nigra ⁄ 10 24 6
Right Substantia Nigra ⁄ 14 20 4 RMGF (BA 10), RFG (BA 19), the RPosG (Ba 40), and the LMTG
(BA 21/39). Subcortical activation was found in the LACC (BA 32),
Occipital Left Lingual Gyrus 18 20 56 2
the left hippocampus, and the RPG (Ba 35). Cerebellar activation
Sub- Left Insula 46 46 18 26
was found in the left declive and left uvula) (Tables 2 and 7, Fig. 1).
lobar Left Amygdala 47 34 24 2
Left Insula 47 46 22 4
Table 5
Temporal Left Fusiform Gyrus-2 37 38 56 12
Disgust-related ALE results organized by Brodmann area.
Left Superior Temporal Gyrus 38 48 14 16
Right Fusiform Gyrus 37 38 52 16 Disgust
Right Superior Temporal Gyrus 41 52 18 6
Lobe Region BA x y z
Anterior Left Culmen ⁄ 26 48 14
Frontal Left Inferior Frontal 47 40 22 14
Table 4 Gyrus-1
Anxiety-related ALE results organized by Brodmann area. Right Inferior Frontal 9 44 12 22
Gyrus-1,2
Anxiety Right Cingulate Gyrus 32 4 18 34
Right Middle Frontal 46 46 20 22
Lobe Region BA x y z
Gyrus
Frontal Right Inferior Frontal 9 38 6 26 Right Inferior Frontal 47 40 20 10
Gyrus-1 Gyrus-4
Limbic Left Anterior Cingulate ⁄ 12 40 2 Limbic Right 28 16 4 10
Left Parahippocampal ⁄ 18 2 10 Parahippocampal
Gyrus Gyrus-1
Left Cingulate Gyrus 31 10 28 38
Occipital Right Inferior ⁄ 46 72 0
Left Cingulate Gyrus 32 6 18 28
Temporal Gyrus
Right Cingulate Gyrus 31 14 34 38
Right Anterior 32 14 42 6 Parietal Left Postcentral Gyrus 3 40 22 48
Cingulate
Sub- Left Claustrum ⁄ 36 6 8
Midbrain Left Red Nucleus ⁄ 0 14 4 lobar Left Claustrum ⁄ 36 0 8
Left Thalamus ⁄ 6 8 2
Parietal Left Postcentral Gyrus 40 52 28 20
Left Thalamus Lateral Posterior 14 18 12
Sub- Left Claustrum ⁄ 34 10 2 Nucleus
lobar Left Caudate Caudate Head 8 4 0 Left Thalamus Medial Dorsal 2 18 4
Right Thalamus Medial Dorsal 6 18 6 Nucleus
Nucleus Left Insula 13 36 22 0
Right Thalamus Pulvinar 2 28 0 Right Thalamus Mammillary Body 10 16 0
Left Insula 13 34 20 0 Right Lentiform Putamen 24 4 2
Right Insula 13 30 16 6 Nucleus
Right Insula 13 36 16 4 Right Insula 13 34 20 2
Temporal Left Superior Temporal 38 44 8 8 Temporal Left Fusiform Gyrus-2 37 42 60 10
Gyrus Right Fusiform Gyrus 37 42 58 12

Please cite this article in press as: Kirby, L. A. J., & Robinson, J. L. Affective mapping: An activation likelihood estimation (ALE) meta-analysis. Brain and
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6 L.A.J. Kirby, J.L. Robinson / Brain and Cognition xxx (2015) xxx–xxx

Table 6 Table 7
Fear-related ALE results organized by Brodmann area. Happiness-related ALE results organized by Brodmann area.

Fear Happiness
Lobe Region BA x y z Lobe Region BA x y z
Anterior Right Culmen ⁄ 38 50 18 Anterior Left Culmen ⁄ 36 44 20
Frontal Right Inferior Frontal 9 46 12 28 Frontal Left Inferior Frontal Gyrus 47 30 20 8
Gyrus-2 Right Middle Frontal Gyrus 10 38 38 16
Left Middle Frontal 46 44 18 22 Right Inferior Frontal 44 54 6 12
Gyrus Gyrus-3
Left Inferior Frontal 47 46 14 0 Right Inferior Frontal Gyrus 47 42 16 10
Gyrus-2 Right Inferior Frontal 47 46 24 6
Gyrus-5,6
Limbic Right Parahippocampal Amygdala 20 4 12
Gyrus-1,2 Limbic Left Parahippocampal Amygdala 22 6 12
Left Anterior Cingulate ⁄ 12 38 0 Gyrus
Left Parahippocampal Amygdala 20 4 10 Left Anterior Cingulate Hippocampus 32 24 14
Gyrus Left Anterior Cingulate 32 0 40 8
Left Anterior Cingulate 24 2 26 4 Right Parahippocampal Amygdala 24 2 12
Left Anterior Cingulate 28 18 24 8 Gyrus-2
Left Anterior Cingulate 32 8 42 10 Right Anterior Cingulate 32 2 48 2
Left Cingulate Gyrus 32 2 22 28 Right Parahippocampal 35 16 28 10
Right Parahippocampal Hippocampus 36 16 14 Gyrus
Gyrus Right Parahippocampal 35 20 20 8
Right Cingulate Gyrus 32 2 8 40 Gyrus
Right Anterior Cingulate 32 4 46 4
Occipital Right Fusiform Gyrus 19 40 74 10
Midbrain Left Red Nucleus ⁄ 2 26 14
Parietal Right Postcentral Gyrus 40 58 28 22
Right Mammillary Body ⁄ 2 14 6
Posterior Left Tuber ⁄ 44 64 26
Occipital Right Inferior Temporal ⁄ 46 70 0
Left Declive ⁄ 38 56 16
Gyrus
Right Declive ⁄ 22 60 12
Right Fusiform Gyrus 19 30 82 14
Right Declive ⁄ 32 80 16
Posterior Left Declive ⁄ 32 68 16 Right Declive ⁄ 28 56 18
Right Declive ⁄ 32 64 20
Temporal Left Middle Temporal 21 52 4 16
Left Insula 13 30 22 0
Gyrus
Right Insula ⁄ 38 16 2
Left Middle Temporal 21 52 4 14
Right Thalamus Medial Dorsal 4 18 10
Gyrus
Nucleus
Left Fusiform Gyrus-1 37 44 50 16
Right Thalamus Pulvinar 2 30 0
Left Superior Temporal 38 48 12 18
Temporal Left Fusiform Gyrus-1 37 38 46 18 Gyrus
Left Fusiform Gyrus 1, 2 37 40 54 16 Left Middle Temporal 39 42 70 16
Left Superior Temporal 42 54 30 18 Gyrus
Gyrus

3.1.6. Humor
The humor search yielded (5 papers, 98 subjects, 10 experi-
ments, 10 conditions, and 79 locations. No cortical or subcortical
activation unique to humor was found. Cerebellar activation was
found in the right culmen (Tables 2 and 8, Fig. 1).
3.1.7. Sadness
The sadness search yielded 54 papers, 1078 subjects, 139 exper-
iments, 161 conditions, and 840 locations. Cortical activation
unique to sadness was found in the RIFG (BA 45), the left middle
frontal gyrus (LMidFG; BA9/47) the left medial frontal gyrus
(LMedFG; BA 9), the right precentral gyrus (RPreG; BA 44), and
the bilateral FG. Subcortical activation was found in the LACC (Ba
32), the right insula (BA 13), the left medial globus pallidus
(LMGP), the right putamen, and the left thalamus (nucleus unspec-
ified) (Tables 2 and 9, Fig. 1).
3.2. Convergent networks
3.2.1. Cortical activation
Most notably, as shown in Table 1, all emotions showed activa-
tion in the dorsolateral prefrontal cortex (DLPFC; BA 9/46, 44, and
47) in the right inferior frontal gyrus (RIFG). The left inferior frontal
gyrus (LIFG; BA 47) demonstrated consistent activation for anger,
disgust, and fear studies. The left middle frontal gyrus (LMFG; BA
9/46) was associated with anger, fear, and sadness. Other notable
areas of convergent activation among different emotion categories
included the fusiform gyrus (FG; BA 37) associated in the left with
fear, and happiness, and with anger and disgust bilaterally; and the
left superior temporal gyrus (LSTG; BA 38) with anger and happi-
ness, and the right STG with disgust and fear.
3.2.2. Limbic/subcortical activation
We observed much expected limbic and other sub-cortical
activity. Specifically, the left cingulate (LCG; BA 32) activated con-
sistently in fear and anxiety studies. The anterior cingulate cortex
(ACC; BA 32) was activated bilaterally during fear studies, with
anxiety on the left and happiness on the right. The amygdala and
parahippocampal gyrus (PG; BA 28) were associated with anxiety
(left), disgust (right), fear (bilateral), happiness (bilateral), and sad-
ness (right). Other subcortical activation included the left caudate
in anxiety and sadness, the left claustrum in anxiety and disgust,
and bilateral insula (BA13) in disgust, anxiety, and fear. The left lat-
eral posterior nucleus (LLPN) of the thalamus showed activation
associated with disgust, the left medial dorsal nucleus (LMDN)
with fear, and both the right pulvinar and MDN with anxiety and
fear.
3.2.3. Cerebellum
Interestingly, there was also convergent activation in the cere-
bellum, with humor and happiness associated with activity in
the left culmen in the anterior portions of the cerebellum, while
the posterior portions were associated with happiness (right
declive and left tuber) and humor (left tuber).

Please cite this article in press as: Kirby, L. A. J., & Robinson, J. L. Affective mapping: An activation likelihood estimation (ALE) meta-analysis. Brain and
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 L.A.J. Kirby, J.L. Robinson / Brain and Cognition xxx (2015) xxx–xxx 7

Fig. 1. Neural networks associated with the seven emotion categories of the BrainMap database. Maps are the result of activation likelihood estimation (ALE) meta-analysis
thresholded at p < 0.01 with a minimum cluster size of 200 mm3.

Table 8 4. Discussion
Humor-related ALE results organized by Brodmann area.
Using ALE on neuroimaging articles in the BrainMap database,
Humor
we found activation in the cortex, subcortical structures, and the
Lobe Region BA x y z
cerebellum. Since there were distinct networks found for individ-
Anterior Left Culmen ⁄ 38 38 26 ual emotions as well as overlap between some emotions, the
Right Culmen ⁄ 26 56 20
results of our data-driven approach most strongly support a
Right Culmen ⁄ 26 42 16
multi-system model of emotions, namely, the affect programs
Cerebellum⁄ Right Cerebellum⁄ ⁄ 4 76 32
theory.
Frontal Right Inferior Frontal ⁄ 46 28 8 All emotion categories queried appear to be represented in the
Gyrus-6
DLPFC (BA 9/46, 44/45, and 47). BA 9 and 46 have been linked to
Limbic Left Parahippocampal Amygdala 18 4 14 processing emotions (Bermpohl et al., 2006) and making decisions
Gyrus
regarding emotions and reward (Deppe, Schwindt, Kugel,
Posterior Left Tuber ⁄ 46 64 24 Plassmann, & Kenning, 2005; Rogers et al., 1999). We also found
Left Uvula ⁄ 30 72 24
activation in Broca’s area (BA 44/45). The right Broca’s area has
been implicated in emotional expression and production of affec-
tive prosody (Wildgruber et al., 2005). In our study this may indi-
cate a role of BA 44/45 in selective attention to certain emotional
Table 9 stimuli such as that contained in speech. However, we were not
Sadness-related ALE results organized by Brodmann area.
able to rule out the possibility of this activation being
Sadness task-dependent (e.g., related to stimuli with verbal content) with-
Lobe Region BA x y z out further investigation.
The involvement of BA 47 across multiple emotions may also
Frontal Left Medial Frontal Gyrus 9 4 44 28
Left Medial Frontal Gyrus 9 6 28 28 indicate affective components of language processing (Sahin,
Left Superior Frontal Gyrus 9 8 52 24 Pinker, & Halgren, 2006; Wildgruber et al., 2005), or could be
Left Middle Frontal Gyrus 9 36 22 32 related to episodic memory (Ranganath, Johnson, & D’Esposito,
Left Middle Frontal Gyrus 9 42 12 28 2003). Our findings of prefrontal emotional convergence seem
Left Middle Frontal Gyrus 47 40 34 6
reflective of an interaction of higher cognitive and affective func-
Right Middle Frontal Gyrus 9 44 8 38
Right Precentral Gyrus 44 50 14 6 tions, potentially highlighting its role as a hub for the integration
Right Inferior Frontal Gyrus 45 46 24 16 of these processes. Notably, these areas have been successfully tar-
Right Inferior Frontal Gyrus-4 47 38 20 10 geted with repeated transcranial magnetic stimulation (rTMS) to
Limbic Left Anterior Cingulate 32 8 18 10 treat otherwise-intractable depression (Isenberg et al., 2005;
Right Parahippocampal Amygdala 26 4 16 Menkes, Bodnar, Ballesteros, & Swenson, 1999).
Gyrus-2
As expected, in contrast to more cognitive processing of emo-
Right Parahippocampal 28 14 4 10
Gyrus-1 tions, our findings also support a more primitive subcortical com-
ponent to emotional processing, especially where greater numbers
Sub- Left Thalamus ⁄ 0 4 8
lobar Left Caudate Caudate Head 2 6 2 of negatively-valenced emotions showed activation (Tables 3–5).
Left Lentiform Nucleus Medial 16 2 8 The insula was predictably activated, as it has previously been
Globus implicated in as broad of functions as autonomic arousal associ-
Pallidus ated with emotions (Kuniecki, Urbanik, Sobiecka, Kozub, &
Right Lentiform Nucleus Putamen 18 4 8
Binder, 2003) and making emotional judgments (Gorno-Tempini
Right Insula 13 48 12 2
et al., 2001). Amygdala, hippocampal, basal ganglia and thalamic

Please cite this article in press as: Kirby, L. A. J., & Robinson, J. L. Affective mapping: An activation likelihood estimation (ALE) meta-analysis. Brain and
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8 L.A.J. Kirby, J.L. Robinson / Brain and Cognition xxx (2015) xxx–xxx

activity were expected limbic components as well, as proposed by
Maclean (1949). Such results are in line with our expectations
demonstrate the robustness of ALE to address this research ques-
tion and similar ones.
We found a small amount of cerebellar activation associated
only with happiness and humor. There were only fives studies
coded for humor, leaving open the possibility that the small cere-
bellar activation we found here was spurious or dependent on the
tasks used. The results associated with happiness have more of a
foundation, however. The role of the cerebellum in motor func-
tions and procedural memory has long been studied (e.g.,
Manto et al., 2012; Molinari et al., 1997; Pascual-Leone et al.,
1993), but its role in emotions has only much more recently been
acknowledged (cite: see Turner et al., 2007). Particularly, Turner
and colleagues found reduced positive affect to typically
positive-affective stimuli in patients with cerebellar lesions. The
revelation of the understudied happiness-related activation in
the cerebellum in our study points further to the robustness of
the ALE method.
Anger had very little common activation (Table 3, Fig. 1), which
could point to a few distinct characteristics and issues. The dearth
of common activation could indicate greater differences across
studies in the elicitation of anger than for other emotions. Anger
is particularly difficult to elicit in the laboratory without very
high-intensity stimuli and often deceit (Harmon-Jones, Amodio,
& Zinner, 2007). Alternatively, the dearth of common activation
in our study may be due to individual differences in the experience
and expression of anger. For example, there is contention as to
whether to categorize anger as an approach or a withdrawal emo-
tion, particularly because there are differences between individuals
on that motivational dimension (Harmon-Jones, 2003). Even for
those who accept anger as an approach-motivating emotion (e.g.,
Carver & Harmon-Jones, 2009), anger still poses a problem. Most
emotions’ approach/withdrawal classifications follow the posi-
tive–negative affect dichotomy – that is, ‘‘positive’’ emotions moti-
vate us to approach an appetitive stimulus, and ‘‘negative’’
emotions motivate organisms to withdraw from an aversive stim-
ulus. Anger is uniquely high-arousal and negatively-valenced,
while motivating many organisms to approach an aversive stimu-
lus (e.g., to fight it). However, the activation we found uniquely
related to anger, with the exception of two coordinates, fell in
the left hemisphere, consistent the long-held position that
approach-related emotions primarily activate in the left hemi-
sphere (e.g., Fetterman, Ode, & Robinson, 2013; Kinsbourne, 1978).
All activation unique to anxiety (Tables 2 and 4; Figs. 2, 3 and 4)
was right-lateralized. Placing anxiety within the approach-
withdrawal framework classifies anxiety as a withdrawal emotion.
Withdrawal emotions are predicted to activate primarily in the
right hemisphere, which we found in the case of anxiety. Further
testing for differential hemispheric lateralization across the
approach-withdrawal distinction yielded no evidence for such a
hypothesis for disgust, fear, happiness, humor, or sadness.
The disgust-related activation was particularly unique (Figs. 2
and 5). Strikingly absent from these metadata were pockets of
insula activity normally characterizing disgust, as has been found
in previous studies (e.g., Phillips et al., 1997). Instead more cingu-
late and DLPFC activation was found, both consistently activated
across many emotional categories. One interesting result was a
point of activation in the left postcentral gyrus. Disgust-related
activation in the somatosensory cortex could reflect disgust’s
adaptive somatosensory-based function as motivation for avoiding
contamination or mutilation.
Other networks specific to affective categories investigated con-
firm much of what is already known about fear and sadness. Fear

Fig. 2. The right inferior frontal gyrus indicated in 3D representations of (a) anger, (b) anxiety, (c) disgust, (d) fear, (e) happiness, (f) humor, and (g) sadness networks. Panels
h, i, and j depict the region of convergence (shown in white) in sagittal, coronal, and axial planes, respectively.

Please cite this article in press as: Kirby, L. A. J., & Robinson, J. L. Affective mapping: An activation likelihood estimation (ALE) meta-analysis. Brain and
Cognition (2015), http://dx.doi.org/10.1016/j.bandc.2015.04.006
 L.A.J. Kirby, J.L. Robinson / Brain and Cognition xxx (2015) xxx–xxx 9

activation (Fig. 2 and Table 6) was located primarily in the fusiform
and superior temporal gyri, the former associated with responses
to emotion in visual stimuli (Lane, Chua, & Dolan, 1999; Ushida
et al., 2008), while the latter contains mirror neurons (Calvert &
Campbell, 2003; Pekkola et al., 2005). Cingulate and amygdala acti-
vation could point to the limbic role in attention to threat stimuli
(e.g., review in Lindquist et al., 2012). Sadness (Fig. 2 and
Table 9), seems to have a strong prefrontal basis as compared to
other emotions. The interpretation specificity of sadness being so
located in the DLPFC is unclear, as the area is associated with emo-
tional decision-making in general.
Our results suggest a multi-system model of affect programs
(basic emotions; Ekman, 1999) as found by Murphy et al. (2003),
supported by the strong quantitative meta-analytic technique
used. However, we did not replicate their findings of fear being
specialized to the amygdala, disgust to the insula and globus

Fig. 3. Negative emotions in sagittal view.

Please cite this article in press as: Kirby, L. A. J., & Robinson, J. L. Affective mapping: An activation likelihood estimation (ALE) meta-analysis. Brain and
Cognition (2015), http://dx.doi.org/10.1016/j.bandc.2015.04.006
10 L.A.J. Kirby, J.L. Robinson / Brain and Cognition xxx (2015) xxx–xxx

pallidus, and anger to the lateral orbitofrontal cortex (OFC).
Instead, our results indicate that these affective categories engage
a more distributed network including many limbic structures, in
line with how anger can serve to prepare the body for
fight-or-flight functions, as well as much pre-frontal involvement,
as we found with most emotion categories investigated. Similarly,
we did not find support for disgust being centralized to the insula,
but instead found it distributed with more prefrontal and
post-central gyrus involvement, as well as subcortical components
in the putamen, mammillary body, and thalamic nuclei. We found
fear to have some amygdalar involvement as they did, but also
throughout fusiform and temporal areas, as well as the cingulate,
hippocampus and cerebellum. Lindquist et al. (2012) eschewed a
locationist viewpoint, contradicting an affect programs model
due to lack of consistency and specificity within emotion cate-
gories across studies. However, they did not show strong support
for a constructionist viewpoint, but only entertained the notion
due to lack of locationist evidence. Our investigation is focused
in what Lindquist and colleagues might call a locationist fashion.
We conceptually replicated the finding of evidence for a
multi-system model of emotions like Murphy et al. (2003) and
Vytal and Hamann (2010), and found more distributed networks
than the former, making ‘‘locationism’’ a weak term to describe
our investigation of emotion networks.
We replicated Vytal and Hamann’s (2010) consistency and
specificity across affect programs using activation likelihood esti-
mation (ALE). Their conclusions are bolstered by our use of a rigor-
ously coded database for neuroimaging papers, and the robustness
of ALE as a meta-analytic technique. Our failure to replicate the
work of either Murphy et al. (2003) or Lindquist et al. (2012) could
be due to using different selection criteria and meta-analytic tech-
niques. Our investigation replicates and bolsters prefrontal their

Fig. 4. Negative emotions in coronal view.

Please cite this article in press as: Kirby, L. A. J., & Robinson, J. L. Affective mapping: An activation likelihood estimation (ALE) meta-analysis. Brain and
Cognition (2015), http://dx.doi.org/10.1016/j.bandc.2015.04.006
 L.A.J. Kirby, J.L. Robinson / Brain and Cognition xxx (2015) xxx–xxx 11

Fig. 5. Negative emotions in axial view.

results with more papers selected from a rigorously coded data-
base using the same robust statistical meta-analytic method.
Together, our results support a strong, right-lateralized pre-
frontal basis and underlying limbic activity surrounding emotional
processing in general. This makes sense in a clinical framework
when considering, as mentioned above, the use of rTMS to treat
depression (Isenberg et al., 2005; Menkes et al., 1999). The right
prefrontal cortex was associated in our study with all emotions
investigated and suggests a role of this area in emotions in general.
Perhaps stimulation of the area relieves severe depression by
restoring function of emotion in general. In addition to finding
areas of convergence among emotions, we found networks unique
to some affective categories. These specific emotions show evi-
dence for distinct neural patterns or affect programs unique to
these basic emotions investigated.
4.1. Limitations
Although our study provides a strong foundation for models of
affect-specific and affect-convergent networks, we acknowledge
several limitations which should be addressed in future studies.
First, we limited our literature search to studies included in the
BrainMap database, which has been estimated to contain around
20% of all neuroimaging papers (Derrfuss & Mar, 2009). Getting
more extensive coverage of the neuroimaging studies on emotion
would be ideal. The database continues to grow and allows for
researchers to rigorously code the studies they wish to see in the
database using the Scribe tool. As the database acquires depth,
affective maps will become increasingly robust and such searches
should become more representative of neuroimaging literature at
large. This will enable more confident neural network maps of

Please cite this article in press as: Kirby, L. A. J., & Robinson, J. L. Affective mapping: An activation likelihood estimation (ALE) meta-analysis. Brain and
Cognition (2015), http://dx.doi.org/10.1016/j.bandc.2015.04.006
12 L.A.J. Kirby, J.L. Robinson / Brain and Cognition xxx (2015) xxx–xxx
less-prominently studied emotions, such as humor. The studies we
queried used various emotion elicitation techniques, which are
beyond the scope of this paper. An increased search set would
allow for the effects of task (visual versus auditory, for example)
to be parsed, which would allow for greater specificity when it
comes to affect maps. However, ALE pools activations, meaning
activations found from ALE reflect coordinates commonly activated
across all studies queried from the database. Therefore, the activa-
tion likely does not vary across emotion elicitation techniques. I
addition to parsing an increased search set by elicitation technique,
future work of ours focuses on improving emotion elicitation tech-
niques by creating stronger stimulus sets using affective film clips.
4.2. Future directions
Our research provides initial models of affective maps that
could be used in future advanced analyses. Specifically this meta-
data would lend itself to a rigorous test of the right-hemisphere
hypothesis and the dual-system models of emotion, particularly
the positive–negative valence distinction. Connectivity analyses
between the significant areas of activation that we have identified,
such as the DLPFC and subcortical structures like the amygdala and
hippocampus, may yield new insights into affective processing dif-
ferences between populations.
5. Conclusions
Using activation likelihood estimation to analyze neuroimaging
metadata on emotion provides the opportunity to create neuro-
functional profiles of each of the basic emotions. These profiles
can be used to help validate and interpret future neuroimaging
work, or provide data-drive a priori selection of brain regions to
include in advanced imaging analyses such as effective connectiv-
ity modeling. Improved emotion profiles could potentially have
particularly relevant applications in clinical work, such as in diag-
nosis and management of psychopathology. While we provide a
foundation for these types of models, we also highlight the need
for increased depth of community database recourses.
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