Hostname: page-component-7c8c6479df-5xszh Total loading time: 0 Render date: 2024-03-28T07:54:45.739Z Has data issue: false hasContentIssue false

The brain basis of emotion: A meta-analytic review

Published online by Cambridge University Press:  23 May 2012

Kristen A. Lindquist
Affiliation:
Department of Neurology, Harvard Medical School/Massachusetts General Hospital/Martinos Center for Biomedical Imaging, Charlestown, MA 02129, and Department of Psychology, Harvard University, Cambridge, MA 02138. lindqukr@nmr.mgh.harvard.eduhttp://www.nmr.mgh.harvard.edu/~lindqukr/
Tor D. Wager
Affiliation:
Department of Psychology and Neuroscience, University of Colorado, Boulder, CO 80309. tor.wager@colorado.eduhttp://www.psych.colorado.edu/~tor/
Hedy Kober
Affiliation:
Department of Psychiatry, Yale University School of Medicine, New Haven, CT 06519. hedy.kober@yale.eduhttp://medicine.yale.edu/psychiatry/people/hedy_kober.profile
Eliza Bliss-Moreau
Affiliation:
California National Primate Research Center, University of California, Davis, CA 95616, and Department of Psychiatry and Behavioral Sciences, University of California, Davis, CA 95616. eblissmoreau@ucdavis.eduhttp://www.elizablissmoreau.com/EBM/home.html
Lisa Feldman Barrett
Affiliation:
Department of Psychology, Northeastern University, Boston, MA 02115, and Departments of Radiology and Psychiatry, Harvard Medical School/Massachusetts General Hospital/Martinos Center for Biomedical Imaging, Charlestown, MA 02129. l.barrett@neu.eduhttp://www.affective-science.org/
Rights & Permissions [Opens in a new window]

Abstract

Researchers have wondered how the brain creates emotions since the early days of psychological science. With a surge of studies in affective neuroscience in recent decades, scientists are poised to answer this question. In this target article, we present a meta-analytic summary of the neuroimaging literature on human emotion. We compare the locationist approach (i.e., the hypothesis that discrete emotion categories consistently and specifically correspond to distinct brain regions) with the psychological constructionist approach (i.e., the hypothesis that discrete emotion categories are constructed of more general brain networks not specific to those categories) to better understand the brain basis of emotion. We review both locationist and psychological constructionist hypotheses of brain–emotion correspondence and report meta-analytic findings bearing on these hypotheses. Overall, we found little evidence that discrete emotion categories can be consistently and specifically localized to distinct brain regions. Instead, we found evidence that is consistent with a psychological constructionist approach to the mind: A set of interacting brain regions commonly involved in basic psychological operations of both an emotional and non-emotional nature are active during emotion experience and perception across a range of discrete emotion categories.

Type
Target Article
Copyright
Copyright © Cambridge University Press 2012

1. Introduction

William James framed the question of emotion–brain correspondence when he wrote, “of two things concerning the emotions, one must be true. Either separate and special centres, affected to them alone, are their brain-seat, or else they correspond to processes occurring in the motor and sensory centres already assigned” (James Reference James1890/1998, p. 473). In this target article, we statistically summarize the last 15 years of neuroimaging research on emotion in an attempt to determine which of these alternatives is correct. We examine the utility of two different models of emotion that have each existed since the beginning of psychology.

2. A locationist account of the brain basis of emotion

A locationist account of emotion assumes that the category emotion and individual categories such as anger, disgust, fear, happiness, sadness (and perhaps a few others) are respected by the body and brain (see Barrett [2006a] for a discussion). The guiding hypothesis of this natural kind model (Barrett Reference Barrett2006a) or modal model (Barrett et al. Reference Barrett, Ochsner, Gross and Bargh2007d) of emotion is that different emotion categories refer to states with endowed motivational characteristics that drive cognition and behavior. It is assumed that these states are biologically basic and inherited, and cannot be broken down into more basic psychological components (Ekman & Cordaro Reference Ekman and Cordaro2011; Izard Reference Izard2011; Panksepp & Watt Reference Panksepp and Watt2011). Despite these common assumptions, there is variability in how different researchers define emotions as natural kinds. Some theorists emphasize the universal characteristics of emotion categories, suggesting that each emotion category (e.g., anger) refers to a “family” of states that share a distinctive universal signal (e.g., facial behavior), physiology, antecedent events, subjective experience, and accompanying thoughts and memories (e.g., Ekman & Cordaro Reference Ekman and Cordaro2011). In this view, emotions can be shaped by culture and learning, but all humans possess the capacity to experience and perceive the same core set of emotion categories.

Other theorists take a developmental approach and argue that all infants are born with a set of “first order emotions” that are evolutionarily given reactions (including feelings, motivations and behaviors) to specific stimuli (e.g., Izard Reference Izard2011). First order emotions form the core of the more elaborate “emotion schemas” that develop with age and learning and consist of complex combinations of emotions, cognitions, and behaviors. Still other theorists emphasize the evolutionary aspect of emotion categories, and argue that emotions are specific behavioral adaptations that are shared with other mammalian species and passed down through phylogeny (e.g., Panksepp Reference Panksepp1998; Reference Panksepp2007; Panksepp & Watt Reference Panksepp and Watt2011). Some models taking an “appraisal” approach to emotion also draw on natural kind assumptions about emotions (cf. Barrett Reference Barrett2006a) by hypothesizing that dedicated cognitive mechanisms automatically make meaning of a stimulus and trigger the corresponding discrete emotion (e.g., Roseman Reference Roseman1984; Ellsworth & Scherer Reference Ellsworth, Scherer, Davidson, Scherer and Goldsmith2003). Relatively little work from an appraisal perspective has investigated the brain basis of emotion (although see Sander et al. Reference Sander, Grafman and Zalla2003; Reference Sander, Grandjean, Kaiser, Wehrle and Scherer2007). Therefore, we do not discuss appraisal models further in this article.

All natural kind models share the assumption that different emotion categories have their roots in distinct mechanisms in the brain and body. The mechanisms underlying discrete emotion categories have been discussed as residing within particular gross anatomical locations (e.g., Calder Reference Calder2003; Ekman Reference Ekman, Dalgleish and Powers1999) or networks (e.g., Izard Reference Izard2011; Panksepp Reference Panksepp1998) in the brain. These models constitute a locationist account of emotion because they hypothesize that all mental states belonging to the same emotion category (e.g., fear) are produced by activity that is consistently and specifically associated with an architecturally defined brain locale (see sections 5.1–5.4 further on)Footnote 1 or anatomically defined networks of locales that are inherited and shared with other mammalian species (Panksepp Reference Panksepp1998; Panksepp & Watt Reference Panksepp and Watt2011). Not all natural kind models are locationist, however; for example, some models propose that each discrete emotion is triggered by an inherited mechanism (e.g., an “affect program”; Ekman & Cordaro Reference Ekman and Cordaro2011; Tomkins Reference Tomkins1962; Reference Tomkins1963) that does not necessarily correspond to a particular brain locale but rather to a specific pattern of autonomic nervous system activity. Much of the contemporary research on emotion makes locationist assumptions; in this article we focus on the models that hypothesize single brain regions to be consistently and specifically associated with different emotion categories, because they represent the most frequent hypothesis that has been tested in the cognitive neuroscience literature. We discuss specific predictions of the locationist approach in section 5, Testing Hypotheses of Brain–Emotion Correspondence (also see Fig. 1).

Figure 1. Locationist Hypotheses of Brain–Emotion Correspondence. A: Lateral view. B: Sagital view at the midline. C: Ventral view. D: Coronal view. Brain regions hypothesized to be associated with emotion categories are depicted. Here we depict the most popular locationist hypotheses, although other locationist hypotheses of brain–emotion correspondence exist (e.g., Panksepp, Reference Panksepp1998). Fear: amygdala (yellow); Disgust: insula (green); Anger: OFC (rust); Sadness: ACC (blue). A color version of this image can be viewed in the online version of this target article at http://www.journals.cambridge.org/bbs.

3. A psychological constructionist account of the brain basis of emotion

A psychological constructionist account of emotion assumes that emotions are psychological events that emerge out of more basic psychological operations that are not specific to emotion. In this view, mental categories such as anger, sadness, fear, et cetera, are not respected by the brain (nor are emotion, perception, or cognition, for that matter; Barrett Reference Barrett2009a; Duncan & Barrett Reference Duncan and Barrett2007; Pessoa Reference Pessoa2008). A psychological constructionist approach to emotion is as old as the locationist approach, at least in its nascent form (e.g., Wundt, James, and other early psychologists were psychological constructionists; see Gendron & Barrett Reference Gendron and Barrett2009). Our contemporary psychological constructionist approach shares much in common with cognitive neuroscience approaches arguing that basic psychological operations are common across diverse task domains (Cole & Schneider Reference Cole and Schneider2007; Dosenbach et al. Reference Dosenbach, Visscher, Palmer, Miezin, Wenger, Kang, Burgund, Grimes, Schlaggar and Petersen2006; Smith et al. Reference Smith, Fox, Miller, Glahn, Fox, Mackay, Filippini, Watkins, Toro, Laird and Beckman2009; van Snellenberg & Wager 2009; Wager et al. Reference Wager, Jonides, Smith and Nichols2005; Wager & Smith Reference Wager and Smith2003). As in the neural context hypothesis, it assumes that the psychological function of individual brain regions is determined, in part, by the network of brain regions it is firing with (McIntosh Reference McIntosh2004). It is also consistent with recent evidence that large-scale networks intrinsic to the brain interact to produce psychological events (Seeley et al. Reference Seeley, Menon, Schatzberg, Keller, Glover, Kenna, Reiss and Greicius2007; Smith et al. Reference Smith, Fox, Miller, Glahn, Fox, Mackay, Filippini, Watkins, Toro, Laird and Beckman2009; Spreng et al. Reference Spreng, Mar and Kim2009; see Deco et al. [2011] for a review). In philosophy of mind, it is consistent with both a token identity and a supervenience approach to mind–brain correspondence (Barrett 2011) and the mental mechanisms approach (Bechtel Reference Bechtel2008). We discuss the psychological constructionist view in a bit more detail because it is unfamiliar to many readers.

In our psychological constructionist model, called the “conceptual act model,” emotions emerge when people make meaning out of sensory input from the body and from the world using knowledge of prior experiences. Emotions are “situated conceptualizations” (cf. Barsalou Reference Barsalou2003) because the emerging meaning is tailored to the immediate environment and prepares the person to respond to sensory input in a way that is tailored to the situation (Barrett Reference Barrett2006b). “Conceptual acts” could also be called “perceptual acts” because they are thought to emerge in consciousness just as visual and auditory percepts do when sensory input is automatically and effortlessly made meaningful using knowledge from prior experience. The idea is that the brain makes an initial prediction about the meaning of the sensory array in context (Bar Reference Bar2007), and the error between this initial top-down prediction and the sensory activity is quickly minimized (Friston Reference Friston2010) to produce a unified conscious field.

In psychological construction, all mental states, whether they are experienced as an instance of a discrete emotion category or not, are realized by more basic psychological operations or “ingredients” of the mind. The goal of psychology is to identify these psychological operations as “psychological primitives,” or the most basic psychological descriptions that cannot be further reduced to anything else mental (because at that point they would describe biological mechanisms). These basic psychological operations are not functionally specific to any discrete emotion category, or even to the category emotion itself. Instead, they are functionally selective for emotion on certain occasions. Because our own model is relatively new, our current model has not identified the most primitive level of psychological description, and it is not yet possible to definitively claim what the most basic psychological operations of the mind are. What we propose is a set of basic domains of psychological function that are a first approximation in the trajectory of a longer research program to identify psychological primitives. These will no doubt be refined as research proceeds in the coming years.

One operation in all psychological constructionist models of emotion is some form of sensory input from the body, such as raw somatic, visceral, vascular, and motor cues (James Reference James1884), arousal (Duffy Reference Duffy1957; Mandler Reference Mandler1975; Reference Mandler1990; Schachter & Singer Reference Schachter and Singer1962), or affect (Harlow & Stagner Reference Harlow and Stagner1932; Hunt Reference Hunt1941; Wundt 1897/1998). In our psychological constructionist view, we refer to this basic psychological domain, as “core affect” (Barrett Reference Barrett2006b; Barrett & Bliss-Moreau Reference Barrett and Bliss-Moreau2009; Russell Reference Russell2003; Russell & Barrett Reference Russell and Barrett1999). In psychology, the word “affect” is used to mean anything that is emotional. Core affect, on the other hand, is a term used to describe the mental representation of bodily changes that are sometimes experienced as feelings of hedonic pleasure and displeasure with some degree of arousal (Barrett & Bliss-Moreau Reference Barrett and Bliss-Moreau2009; Russell Reference Russell2003; Russell & Barrett Reference Russell and Barrett1999). Core affect is realized, in part, by visceral control systems that help organisms deal with motivationally salient stimuli in the environment. A functioning peripheral nervous system is not necessary for a person to experience a core affective state (e.g., Critchley et al. Reference Critchley, Mathias and Dolan2001) as long as they have some prior experiences to provide them with central nervous system representations of bodily states. However, in healthy individuals, core affect is usually accompanied by somatovisceral, kinesthetic, proprioceptive, and neurochemical fluctuations that take place within the core of the body and are represented in the brain. Changes in core affect are a homeostatic barometer–the body's way of representing whether objects in the environment are valuable or not in a given context. The concept of core affect shares much in common with the idea that bodily cues constitute a core ingredient in mental life (e.g., the idea that being embodied is essential to consciousness: Craig Reference Craig2009; Damasio Reference Damasio1999; the idea that feelings are a common currency for valuation of objects in the world: Cabanac Reference Cabanac2002). We assume that core affect is not psychologically meaningful unless it is attached to an object; it is made meaningful via a second basic psychological operation, which we describe next.

All psychological constructionist models include a second basic psychological operation by which internal sensory cues or their associated affective feelings are automatically and effortlessly made meaningful (i.e., experienced as related to or caused by an event or object, usually in the external surroundings). Candidates for this second psychological operation include ideas (Wundt 1897/1998), social affiliation (Schachter & Singer Reference Schachter and Singer1962), attribution (Russell Reference Russell2003), or, as we propose in our model, categorization as situated conceptualization (Barrett Reference Barrett2006b). The process of “conceptualization” (and the other operations that support it, such as executive attention) links perceptions of sensory input from the world with input from the body to create a meaningful psychological moment. In our hypothesis, people automatically make meaning of their core affective state by engaging in a situated conceptualization that links it to an object or event. Conceptualization is the process by which stored representations of prior experiences (i.e., memories, knowledge) are used to make meaning out of sensations in the moment (Wilson-Mendenhall et al. Reference Wilson-Mendenhall, Barrett, Simmons and Barsalou2011). A person can make the situated conceptualization that core affect is a physical symptom (e.g., a racing heart), a simple feeling (e.g., feeling tired or wound up), or an instance of a discrete emotion category (e.g., anger vs. fear). And at other times, core affect is perceived as part of an object itself rather than one's reaction to it. For instance, a food is delicious or distasteful, a painting is beautiful or garish, or a person is nice or nasty. Because we hypothesize that people make meaning of their core affective states in context, experiencing them as a part of an emotion, perception, belief, or judgment, a psychological constructionist account does not simply reduce the category of emotion to positive or negative affect (as is often claimed in summaries of “dimensional models of emotion”; e.g., Fontaine et al. Reference Fontaine, Scherer, Roesch and Ellsworth2007; Keltner et al. Reference Keltner, Ekman, Gonzaga, Beer, Davidson, Scherer and Goldsmith2003). Conceptualization can be said to produce cognitive appraisals realizing emotion (Barrett et al. Reference Barrett, Mesquita, Ochsner and Gross2007c), where such appraisals are descriptions of the features or properties of emotional experience (Clore & Ortony Reference Clore, Ortony, Lewis, Haviland-Jones and Barrett2008). In many appraisal models, the assumption is that the brain contains a series of specific cognitive appraisal mechanisms (e.g., there is a specific mechanism to appraise the novelty of an object, or whether one's goals are blocked) which, when configured into a particular pattern, trigger discrete emotions. In our model, we do not propose any new or unique mental processes that cause emotion; instead, we propose a mechanism (categorization) that has been well documented in the psychological and cognitive neuroscience literature. Categorization (or conceptualization) is a fundamental process in the human brain that functions like a chisel, leading people to attend to certain features in a sensory array and to ignore others. Only some of the wavelengths of light striking our retinas are transformed into seen objects, only some of the changes in air pressure registered in our ears are heard as words or music, and only some bodily changes are experienced as emotion. To categorize something is to render it meaningful. It then becomes possible to make reasonable inferences about that thing, to predict what to do with it, and to communicate our experience of it to others. There are ongoing debates about how categorization works, but the fact that it works is not in question.

Definitions

Natural kind approach. A theoretical framework for understanding the ontology of emotions. The natural kinds approach assumes that emotion categories such as anger, sadness, fear, et cetera, map on to biological categories that are given by the brain and body, and cannot be reduced to more basic psychological parts.

Locationist approach. Many national kind models of emotion conform to a locationist approach in which discrete emotion categories (e.g., anger) are assumed to be consistently and specifically localized to discrete brain locales or anatomical networks.

Psychological constructionist approach. A theoretical framework for understanding the ontology of emotions. The psychological constructionist approach assumes that emotion categories such as anger, sadness, fear, and so forth, are common sense categories whose instances emerge from the combination of more basic psychological operations that are the common ingredients of all mental states.

Core affect. The mental representation of bodily sensations that are sometimes (but not always) experienced as feelings of hedonic pleasure and displeasure with some degree of arousal. Core affect is what allows an organism know if something in the environment has motivational salience (i.e., is good for it, bad for it, approachable, or avoidable). Barring organic abnormality, core affect is accompanied by somatovisceral, kinesthetic, proprioceptive, and neurochemical fluctuations that take place within the core of body and feed back to be represented in the brain.

Conceptualization. The process by which sensations from the body or external world are made meaningful in a given context using representations of prior experience. Conceptualization occurs in a situated fashion (as in “situated conceptualization”; see Barslou etal., in press), drawing on the representations of prior experience that are activated by the present physical and psychological situation.

Executive attention. The process by which some representations are selectively enhanced and others are suppressed. This is also known as “endogenous,” “controlled,” or “goal-based” attention and can be exerted both volitionally and without the conscious experience of volition. Executive attention can shape the activity in other processes such as core affect, conceptualization, or language use. In the case of emotion, executive attention foregrounds certain core affective feelings and exteroceptive sensory sensations in a moment, and guides which situated conceptualizations are brought to bear to make meaning of those sensations in the given context.

Emotion words. The set of words that ground the abstract categories that humans experience and communicate about. In the case of abstract categories like emotions, words are “essence placeholders” that help cohere feelings, behaviors, and facial expressions together as instances of a meaningful category.

Neural reference space. The set of neurons that are probabilistically involved in realizing a class of mental events (such as anger, or even emotion).

Functional selectivity. This occurs when a set of neurons show a consistent increase in activation for one mental state (e.g., anger, disgust, emotion) or basic psychological operation (e.g., categorization, core affect) more so than for others in a given instant. The neurons are not specific to any mental state, although they might be more frequently activated in some than in others. Functional selectivity might occur because a brain region supports a more basic psychological operation that helps to construct a certain mental state (e.g., the amygdala supports detection of salient exteroceptive sensations and is functionally selective for perceptions of fear). A brain area might be functionally selective for one mental state or even one basic psychological operation in one instance, and for another state or operation in another instance (e.g., ventromedial prefrontal cortex helps to realize both core affect and conceptualization).

In our model, categorization in the form of situated conceptualization is realized in a set of brain regions that reconstitutes prior experiences for use in the present. This set of brain regions has also been called the “episodic memory network” (e.g., Vincent et al. Reference Vincent, Snyder, Fox, Shannon, Andrews, Raichle and Buckner2006) or the “default network” (e.g., Raichle et al. Reference Raichle, MacLeod, Snyder, Powers, Gusnard and Shulman2001). It is active when people recall the past (e.g., Buckner & Carroll Reference Buckner and Carroll2007; Schacter et al. Reference Schacter, Addis and Buckner2007; see also, McDermott et al. [Reference McDermott, Szpunar and Christ2009] for a meta-analysis), imagine the future (e.g., Addis et al. Reference Addis, Wong and Schacter2007; see also, Hassabis & Maguire Reference Hassabis and Maguire2009; Moulton & Kosslyn Reference Moulton and Kosslyn2009; Schacter et al. Reference Schacter, Addis and Buckner2007), make context-sensitive predictions about others' thoughts and feelings (e.g., as in theory of mind; Saxe & Kanwisher Reference Saxe and Kanwisher2003; see Mitchell Reference Mitchell2009), or make meaning of exteroceptive sensations (e.g., context-sensitive visual perception; Bar et al. Reference Bar, Kassam, Ghuman, Boshyan, Schmid, Dale, Hämäläinen, Marinkovic, Schacter and Rosen2006; see also, Bar 2009). In emotion, we hypothesize that this psychological operation makes a prediction about what caused core affective changes within one's own body or what caused the affective cues (e.g., facial actions, body postures, or vocal acoustics) in another person, and this prediction occurs in a context-sensitive way (with the result that core affect in context is categorized as an instance of anger, disgust, fear, etc.; Barrett Reference Barrett2006b; Reference Barrett2009b; see also, e.g., Barrett & Kensinger Reference Barrett and Kensinger2010; Lindquist & Barrett Reference Lindquist and Barrett2008a; Wilson-Mendenhall et al. Reference Wilson-Mendenhall, Barrett, Simmons and Barsalou2011). When making meaning out of core affect, conceptualization draws on prior experiences and perceptions of emotion to realize the emotional gestalts that make up part of what Edelman calls “the remembered present” (cf. Edelman Reference Edelman1989; see Barrett et al. Reference Barrett, Mesquita, Ochsner and Gross2007c; Barrett Reference Barrett2009b).

Our model proposes two additional operations that are important to the psychological construction of emotion. We hypothesize that emotion words that anchor emotion categories work hand in hand with conceptualization (Barrett Reference Barrett2006b; Barrett et al. Reference Barrett, Lindquist and Gendron2007b). Emotion words are essential to our model because we assume that the instances of any emotion category (e.g., anger) that are created from affective feelings don't have strong statistical regularities in the real world or firm natural category boundaries (for a discussion of the empirical evidence, see Barrett Reference Barrett2006a; 2009; Barrett et al. Reference Barrett, Lindquist, Bliss-Moreau, Duncan, Gendron, Mize and Brennan2007a). In our view, emotion categories are abstract categories that are socially constructed (Barrett Reference Barrett2009a). As with all abstract categories, in the absence of strong perceptual statistical regularities within a category, humans use words as the glue that holds the category together (Barsalou & Weimer-Hastings Reference Barsalou, Wiemer-Hastings, Pecher and Zwaan2005). In fact, even infants routinely use the phonological form of words to make conceptual inferences about novel objects that share little perceptual similarity (Dewar & Xu Reference Dewar and Xu2009; Ferry et al. Reference Ferry, Hespos and Waxman2010; Xu Reference Xu2002), and we believe that adults do the same thing. Because words are in part represented via situated simulations of prior experiences (e.g., Simmons et al. Reference Simmons, Hamann, Harenski, Hu and Barsalou2008), we expect emotion words to work together with conceptualization when perceivers make meaning of core affective states.

Executive attention is the fourth operation that is particular to our psychological constructionist approach (Barrett Reference Barrett2009a; Barrett et al. Reference Barrett, Tugade and Engle2004). Executive attention helps direct the combination of other psychological operations to produce an emotional gestalt. At any point in time, the brain is processing information from the body (core affect), information from outside the body (exteroceptive sensory information), and representations of prior experiences (conceptualizations). For instance, many different representations of the past might become active to make meaning of a core affective state. We hypothesize that executive attention helps determine which representations are utilized to make meaning of that state, and which are suppressed (cf. Barrett Reference Barrett2009b; see Barrett et al. Reference Barrett, Tugade and Engle2004, for a discussion). Executive attention can also control which exteroceptive sensory representations are favored for additional processing, or if core affect is consciously represented in awareness. Importantly, executive attention need not be volitional or effortful and can operate well before subjective experience is generated (Barrett et al. Reference Barrett, Tugade and Engle2004). We acknowledge that additional operations are probably important to the construction of emotion and will be incorporated into our model as research accrues.

In the past, most researchers who found brain correlates of emotion assumed that their results were consistent with a locationist approach (e.g., the basic emotion approach) because these were the only models to map psychological states to a biological level of analysis in a way that was linked to evolution. Constructionist hypotheses (which were typically social, rather than psychological) were restricted to the psychological level in a manner divorced from evolution. But this is an accident of history. In fact, there are very clear brain hypotheses that develop from a psychological constructionist view (Barrett Reference Barrett2006b), and our psychological constructionist approach is the first that attempts to map basic psychological operations to brain networks that comprise instances of a psychological category such as emotion, or to the subordinate categories of anger, sadness, fear, disgust, and happiness (see also Barrett Reference Barrett2006a; Reference Barrett2006b; Reference Barrett2009a; Reference Barrett2009b; Barrett et al. Reference Barrett, Lindquist, Bliss-Moreau, Duncan, Gendron, Mize and Brennan2007a; Reference Barrett, Mesquita, Ochsner and Gross2007c; Kober et al. Reference Kober, Barrett, Joseph, Bliss-Moreau, Lindquist and Wager2008). Our hypothesized psychological operations, as they currently stand, are associated with assemblies of neurons within distributed networks (rather than a one-to-one mapping of ingredient to network).  We hypothesize that these networks combine and constrain one another like ingredients in a recipe, influencing and shaping one another in real time according to the principles of constraint satisfaction (Barrett et al. Reference Barrett, Ochsner, Gross and Bargh2007d). With more research, it will be possible to identify the distributed brain networks that are associated with the most basic psychological operations of the mind.

Together, the functional networks that instantiate basic psychological operations during emotion experiences and perceptions form the “neural reference space for discrete emotion.” According to Gerald Edelman (Reference Edelman1989), a “neural reference space” is made up of the neurons that are probabilistically involved in realizing a class of mental events (such as anger, or even emotion).Footnote 2 The functions of distinct brain areas within the neural reference space are best understood within the context of the other brain areas to which they are connected (either anatomically or because of the timing and coordination of neural activity) and in terms of the basic psychological operations they are functionally selective for in a given instance. Unlike a locationist approach, which hypothesizes that a single brain region will be consistently and specifically activated across instances of a single emotion category, a psychological constructionist approach hypothesizes that the same brain areas will be consistently activated across the instances from a range of emotion categories (and, although it is beyond the scope of this article, even in non-emotional states), meaning that that brain region is not specific to any emotion category (or even to emotion per se). We focus on the brain regions that we believe are hubs in the networks corresponding to basic psychological operations and discuss specific predictions in section 5, “Testing Hypotheses of Brain–Emotion Correspondence” (also see Fig. 2).

Figure 2. Psychological Constructionist Hypotheses of Brain–Emotion Correspondence. A: Lateral view. B: Sagital view at the midline. C: Ventral view. D: Coronal view. Brain regions hypothesized to be associated with psychological operations are depicted. In some cases, we present only the key brain regions within networks that have been empirically linked to our hypothesized psychological operations. In instances where the whole brain network is not depicted, we point readers to relevant literature. Core Affect (pink): amygdala, insula, mOFC (Bas 10m, 11m, 13a, b, 14r, c), lOFC (BAs 47, 12, 13l, m, 11l), ACC (Bas, 32, 24, 25), thalamus, hypothalamus, bed nucleus of the stria terminalis, basal forebrain, PAG. Conceptualization (purple): VMPFC (Bas 11, 25, 32, 34), DMPFC (BAs 9, 10p), medial temporal lobe* (hippocampus, entorhinal cortex, parahippocampal cortex), posterior cingulate cortex/retrosplenial area (BA 23, 31). Language (green): VLPFC (Bas 44, 45, 46), anterior temporal lobe (BA 38); for additional regions, see Vigneau et al. (Reference Vigneau, Beaucousin, Hervé, Duffau, Crivello, Houdé, Mazoyer and Tzourio-Mazoyer2006). Executive Attention (orange): DLPFC (BAs 9, 10, 46), VLPFC (BAs 44, 45, 46); for additional regions see Corbetta & Shulman, (Reference Corbetta and Shulman2002); Dosenbach et al. (Reference Dosenbach, Visscher, Palmer, Miezin, Wenger, Kang, Burgund, Grimes, Schlaggar and Petersen2006); Wager et al. (Reference Wager, Reading and Jonides2004). (*this structure is not visible in this view of the brain). A color version of this image can be viewed in the online version of this target article at http://www.journals.cambridge.org/bbs.

4. Meta-analysis of neuroimaging studies on emotion

In this article, we report a meta-analysis of neuroimaging studies on emotion to assess whether the data are more consistent with a locationist or a psychological constructionist account of emotion. In our meta-analysis, strong evidence for a locationist account would be found if instances of an emotion category (e.g., fear) are consistently and specifically associated with increased activity in a brain region (or a set of regions within an anatomically inspired network) across published neuroimaging studies. Consistency refers to the fact that a brain region shows increased activity for every instance of an emotion category (e.g., the amygdala shows increased activity each time a person experiences an instance of the category fear). Specificity refers to the fact that a given brain region is active for instances of one (and only one) emotion category (e.g., the amygdala does not show increased activity when a person is experiencing an instance of anger, disgust, happiness, or sadness). Support for a psychological constructionist view, in contrast, would be found if the same brain region(s) were involved in realizing instances of several emotion categories–and, furthermore, if the brain region(s) are more generally important to realizing a basic psychological operation (e.g., core affect, conceptualization, language, or executive attention). From this perspective, we would not expect instances of any emotion category to be specifically related to increased activation in any single brain region or set of regions. A brain region might be functionally selective for a given emotion category in a given instance, however, because it helps realize a more basic operation that contributes to the emergent state.

In 2005, we began our meta-analytic project to probe the brain basis of emotion. We have since published one chapter (Wager et al. Reference Wager, Barrett, Bliss-Moreau, Lindquist, Duncan, Kober, Joseph, Davidson and Mize2008) and two papers (Barrett et al. Reference Barrett, Mesquita, Ochsner and Gross2007c; Kober et al. Reference Kober, Barrett, Joseph, Bliss-Moreau, Lindquist and Wager2008) reporting our findings for the neuroimaging studies of discrete emotion and affect that came out between 1990 and 2005. Supporting a psychological constructionist approach to emotion, we found that the neural reference space for emotion and affect could be inductively parsed into six distributed functional groups of brain regions (i.e., regions consistently co-activated across studies) using a series of multidimensional scaling and cluster analyses (Kober et al. Reference Kober, Barrett, Joseph, Bliss-Moreau, Lindquist and Wager2008) (See Fig. 3). These functional groups can be mapped to the hypothesized psychological operations that we derived from behavioral data (e.g., Barrett 2006).

Figure 3. Kober et al.'s (2008) Functional Clusters. Kober et al.'s (2008) six functional clusters are consistent with the ingredients hypothesized by our psychological constructionist model. The brain areas making up the “core limbic group” and “lateral paralimbic group” are part of the network that helps to constitute core affect. Aspects of the “medial posterior group” and “medial PFC group” are part of the network involved with conceptualization. Areas in the “cognitive/motor control group” are consistent with the networks supporting language and executive attention. In addition, an “occipital/visual group” was also identified as part of the neural reference space for emotion. Visual cortex has connectivity with areas involved in core affect (e.g., amygdala, orbitofrontal cortex; Amaral & Price Reference Amaral and Price1984; Barrett & Bar Reference Barrett and Bar2009; Pessoa & Adolphs Reference Pessoa and Adolphs2010), and there is growing evidence that a person's core affective state modulates activity in primary visual cortex (Damaraju et al. Reference Damaraju, Huang, Barrett and Pessoa2009). Core affect even shapes aspects of visual perception ranging from contrast sensitivity (Phelps et al. Reference Phelps, Ling and Carrasco2006) to visual awareness (Anderson et al. Reference Anderson, Siegel, Bliss-Moreau and Barrett2011). A color version of this image can be viewed in the online version of this target article at http://www.journals.cambridge.org/bbs.

4.1. Analysis strategy

In the present article, we expanded upon our initial meta-analytic efforts to directly compare the locationist versus the psychological constructionist approach for neuroimaging studies of discrete emotion. A detailed description of our meta-analytic methods and Figure S1 are included in the supplementary materials (available at: http://www.journals.cambridge.org/bbs2012008). In comparing these hypotheses, we are comparing a hypothesis with very specific empirical requirements (i.e., evidence for consistency and specificity in brain–emotion correspondence) to a hypothesis with more flexible empirical requirements (i.e., evidence of multiple operations across multiple categories). Given the popularity of locationist models of emotion, we made analysis choices that favored a clear and unbiased test of the locationist approach, even though it disadvantaged us in testing the full scope and power of the psychological constructionist approach. After updating our database to include papers from 2006 and 2007, we exclusively sampled studies that focused on discrete emotion experiences or perceptions to increase the likelihood that we would find consistent and specific brain localizations corresponding to these categories, should they exist (see Tables S1 and S2 in supplementary materials for details on the inclusion criteria and database; available at: http://www.journals.cambridge.org/bbs2012008). We also conducted a number of statistical analyses with the potential to yield evidence in favor of a locationist account (outlined in the next section).

4.1.1. The neural reference space for discrete emotion

We began by estimating the neural reference space for discrete emotion. This space refers to the brain regions that show a consistent increase in activation for the experience or perception of instances of anger, sadness, fear, disgust, and happiness. A brain region might appear in this space because its activation consistently increases in studies of one discrete emotion category but not others, some categories but not others, or all categories of emotion. Alternatively, a brain region could appear in this space even when it does not consistently have increased activation during any discrete emotion category per se, but because it has consistent increases during instances of the entire category emotion (e.g., the brain region shows consistent increases in activation in some but not all studies of anger experience, anger perception, fear experience, fear perception, and so forth, so that the region is consistently activated across the category emotion, but is not specific to any discrete emotion category). Our derived neural reference space for discrete emotion (Fig. 4; see also Table S3 in supplementary materials, available at: http://www.journals.cambridge.org/bbs2012008) closely resembles the one reported in Kober et al. (Reference Kober, Barrett, Joseph, Bliss-Moreau, Lindquist and Wager2008), even when limiting our analysis to studies of discrete emotion and including papers from 2006–2007. Next, we examined whether any emotion categories were more likely to be associated with increased activity in certain brain areas than in others.

Figure 4. The Neural Reference Space for Discrete Emotion. The neural reference space (phrase coined by Edelman [1989]) is the set of brain regions consistently activated across all studies assessing the experience or perception of anger, disgust, fear, happiness and sadness (i.e. the superordinate category emotion). Brain regions in yellow exceeded the height threshold (p<05) and regions in orange exceeded the most stringent extent-based threshold (p<001). Regions in pink and magenta correspond to lesser extent-based thresholds and are not discussed in this article. Cortex is grey, the brainstem and nucleus accumbens are green, the amygdala is blue and the cerebellum is purple. A color version of this image can be viewed in the online version of this target article at http://www.journals.cambridge.org/bbs.

4.1.2. Density analyses

Within the neural reference space, we first searched over the brain for voxels with more consistent activation (within 10mm) for instances of one emotion category than for all others (e.g., for voxels that reached family-wise error-rate corrected significance in the comparison [fear perception vs. perception of other categories]). This analysis yielded a series of statistical maps reflecting whether each voxel was more frequently activated in studies of each emotion category versus the average of the others, accounting for the different numbers of studies in different categories and the base-rate of background activation across the brain for each emotion category. These analyses are standard for neuroimaging meta-analysis (see Wager et al. Reference Wager, Lindquist and Kaplan2007) and are described in detail in the supplementary materials. The density analyses speak to whether increases in a brain region are consistently associated with the experience or perception of instances of an emotion category. This provides one kind of information about the consistency and specificity of brain activity for particular emotion categories by considering the activity in each region, for each emotion type, relative to background activation levels across the brain.

4.1.3. χFootnote 2 analyses

We next probed the voxels identified in the density analysis further by asking whether there was any absolute difference in the proportion of contrasts activating near those voxels (within 10mm) for each emotion category versus the others. This was accomplished using χFootnote 2 analyses on the contingency table consisting of counts of study contrasts showing activation in or around these voxels compared to study contrasts without such activations for the target emotion category versus other categories. This analysis yielded a series of statistical maps reflecting whether each voxel was more frequently activated in studies of each emotion category versus the average of the others, irrespective of activations elsewhere in the brain.

Both density and χFootnote 2 analyses speak to whether increased activations in a set of voxels that are consistently associated with the experience or perception of instances of an emotion category are also functionally selectiveFootnote 3 for that emotion category. A region that is functionally selective for instances of an emotion category would show voxels that are significant in both the density analysis and χ Footnote 2 analysis. Functional specificity exists if voxels activated selectively for instances of one emotion category also never show increased activity during instances of any other emotion categories. We did not find evidence for functional specificity with respect to any emotion category in our analyses (i.e., every region that was activated for one emotion category was activated for at least one other category). Therefore, our findings only speak to functional selectivity.

4.1.3. Logistic regressions

Finally, in a third set of analyses we used a series of stepwise logistic regressions to ask which emotion categories and experimental methods predicted increased activity in regions of interest. We present the odds ratios for these regressions (in Table S6 in the supplementary materials, available at http://www.journals.cambridge.org/bbs2012008) or the percent increase in odds that a variable predicted either increased activity in a brain area or no increase in a brain area (in Fig. 5).Footnote 4 The logistic regressions speak to both consistency and specificity of increased brain activation. Consistency is observed when any variable significantly predicted increased activity in a given brain area. Specificity is observed when one variable significantly predicted increased activity in a given brain area but all others significantly predicted no increase in activity. If a variable was not a significant predictor, then it is sometimes associated with increased activity, and is sometimes not.

Figure 5. Logistic Regression Findings. Selected results from the logistic regressions are presented (for additional findings, see Table S6 in supplementary materials). Circles with positive values represent a 100% increase in the odds that a variable predicted an increase in activity in that brain area. Circles with negative values represent a 100% increase in the odds that a variable predicted there would not be an increase in activity in that brain area. Legend: Blue lines: left hemisphere; Green lines: right hemisphere. Arrowheads: % change in odds is greater than values represented in this figure. Abbreviations: OFC: orbitofrontal cortex; DLPFC: dorsolateral prefrontal cortex; ATL: anterior temporal lobe; VLPFC: ventrolateral prefrontal cortex; DMPFC: dorsomedial prefrontal cortex; aMCC: anterior mid-cingulate cortex; sAAC: subgenual ACC. A color version of this image can be viewed in the online version of this target article at http://www.journals.cambridge.org/bbs.

5. Testing hypotheses of brain–emotion correspondence

5.1. The amygdala

According to a locationist hypothesis, the amygdala (Fig. 1, yellow) is either the brain locus of fear or is the most important hub in a fear circuit. This amygdala-fear hypothesis was most clearly popularized by behavioral neuroscience work showing that the amygdala (in particular, the central nucleus) supports the cardiovascular changes that occur when rats freeze or startle in response to tones previously paired with shock (called “fear learning”: LeDoux et al. Reference LeDoux, Sakaguchi and Reis1983; Reference LeDoux, Sakaguchi, Iwata and Reis1985; Reference LeDoux, Ciccetti, Xagoraris and Romanski1990; for reviews see Fanselow & Poulous 2005; Fendt & Fanselow Reference Fendt and Fanselow1999; LeDoux Reference LeDoux2007; Öhman Reference Öhman2009; and “fear potentiated startle”: Davis Reference Davis1992; Hitchcock & Davis Reference Hitchcock and Davis1986; Reference Hitchcock and Davis1987; see Davis et al. Reference Davis, Amaral and Winslow2008; Fendt & Fanselow Reference Fendt and Fanselow1999). Electrical stimulation of the amygdala elicits defensive behavior in rats (e.g., retreat; Maskati & Zbrozyna Reference Maskati and Zbrozyna1989) and enhances startle to acoustic stimuli (Rosen & Davis Reference Rosen and Davis1988). The amygdala-fear hypothesis was further strengthened by evidence that humans show increased amygdala activity to neutral tones that have been previously paired with noxious noise blasts (i.e., “fear learning”; LaBar et al. Reference LaBar, Gatenby, Gore, LeDoux and Phelps1998). Individuals with amygdala lesions (LaBar et al. Reference LaBar, LeDoux, Spencer and Phelps1995) or atrophy (Bechara et al. Reference Bechara, Tranel, Damasio, Adolphs, Rockland and Damasio1995) show impaired skin conductance responses during “fear learning” and have difficulty perceiving instances of fear in voices (Brierley et al. Reference Brierley, Medford, Shaw and David2004; Scott et al. Reference Scott, Young, Calder, Hellawell, Aggleton and Johnson1997, but see, Adolphs & Tranel Reference Adolphs and Tranel1999; Anderson & Phelps Reference Anderson and Phelps1998), bodies (Sprengelmeyer et al. Reference Sprengelmeyer, Young, Schroeder, Grossenbacher, Federlein, Büttner and Przuntek1999; but see, Atkinson et al. Reference Atkinson, Heberlein and Adolphs2007), and startled faces with wide eyes (e.g., Adolphs et al. Reference Adolphs, Tranel, Damasio and Damasio1994; Reference Adolphs, Tranel, Damasio and Damasio1995; Reference Adolphs, Tranel, Hamann, Young, Calder, Phelps, Anderson, Lee and Damasio1999; although see, Adolphs et al. Reference Adolphs, Gosselin, Buchanan, Tranel, Schyns and Damasio2005; Tsuchiya et al. Reference Tsuchiya, Moradi, Felsen, Yamazaki and Adolphs2009). An individual with bilateral amygdala lesions failed to report fearful experiences when placed in close contact with snakes, spiders, or when startled (Feinstein et al. Reference Feinstein, Adolphs, Damasio and Tranel2011; although see, Anderson & Phelps Reference Anderson and Phelps2002). Finally, the amygdala is implicated in psychopathology involving the experience of anxiety in humans (for a review, see Damsa et al. Reference Damsa, Kosel and Moussally2009; for a meta-analytic review, see Etkin & Wager Reference Etkin and Wager2007).

According to a psychological constructionist view, the amygdala is part of the distributed network that helps to realize core affect (Fig. 2, panel D, bright pink) because it is involved in signaling whether exteroceptive sensory information is motivationally salient (for similar views see Adolphs Reference Adolphs2008; Reference Adolphs2009; Duncan & Barrett Reference Duncan and Barrett2007; Pessoa Reference Pessoa2010b; Pessoa & Adolphs Reference Pessoa and Adolphs2010; Sander et al. Reference Sander, Grafman and Zalla2003; Whalen Reference Whalen2007; Whalen Reference Whalen1998). The amygdala is most likely to be active when the rest of the brain cannot easily predict what sensations mean, what to do about them, or what value they hold in that context. Salient objects or events influence an organism's body state in a way that can be experienced as core affective feelings (Barrett & Bliss-Moreau Reference Barrett and Bliss-Moreau2009). They can also cause the amygdala to signal to other parts of the brain to sustain processing so that uncertainty about the stimulus can be resolved (Whalen Reference Whalen2007). As a result, affect can be considered a source of attention in the brain (Barrett & Bar Reference Barrett and Bar2009; Duncan & Barrett Reference Duncan and Barrett2007; Pessoa Reference Pessoa2008; Reference Pessoa2010b; Vuilleumier Reference Vuilleumier2005; Vuilleumier & Driver Reference Vuilleumier and Driver2007).

From a psychological constructionist point of view, fear-inducing stimuli might fall into the class of uncertain and therefore salient stimuli, but the amygdala is not specific to the category fear. Consistent with this view, the amygdala is routinely implicated in orienting responses to motivationally relevant stimuli (Holland & Gallagher Reference Holland and Gallagher1999). Novel stimuli (e.g., Blackford et al. Reference Blackford, Buckholtz, Avery and Zald2010; Breiter et al. Reference Breiter, Etcoff, Whalen, Kennedy, Rauch, Buckner, Strauss, Hyman and Rosen1996; Moriguchi et al. Reference Moriguchi, Negreira, Weirerich, Dautoff, Dickerson, Wright and Barrett2010; Schwartz et al. Reference Schwartz, Wright, Shin, Kagan, Whalen, McMullin and Rausch2003; Weierich et al. Reference Weierich, Wright, Negreira, Dickerson and Barrett2010; Wilson & Rolls Reference Wilson and Rolls1993; Wright et al. Reference Wright, Martis, Schwartz, Shin, Fischer, McMullin and Rausch2003; Reference Wright, Wedig, Williams, Rauch and Albert2006; Reference Wright, Negreira, Gold, Britton, Williams and Barrett2008), uncertain stimuli (e.g., Herry et al. Reference Herry, Bach, Esposito, Di Salle, Perrig, Scheffler, Lüthi and Seifritz2007), and unusual stimuli (e.g., Blackford et al. Reference Blackford, Buckholtz, Avery and Zald2010) robustly activate the amygdala and produce cardiovascular responses associated with affective changes (Mendes et al. Reference Mendes, Blascovich, Hunter, Lickel and Jost2007). Amygdala lesions disrupt normal responses to novelty and uncertainty in mammals (e.g. Bliss-Moreau et al. Reference Bliss-Moreau, Toscano, Baumann, Mason and Amaral2010; Burns et al. Reference Burns, Annett, Kelley, Everitt and Robbins1996; Mason et al. Reference Mason, Capitanio, Machado, Mendoza and Amaral2006; Missilin & Ropartz Reference Missilin and Ropartz1981; Nachman & Ashe Reference Nachman and Ashe1974; for reviews, see Knight & Grabowecky 1999; Petrides Reference Petrides2007). Individuals with amygdala lesions do not automatically allocate attention to aversive stimuli (Anderson & Phelps Reference Anderson and Phelps2001) and socially relevant stimuli (Kennedy & Adolphs Reference Kennedy and Adolphs2010), as do individuals with intact amygdalae. Amygdala responses habituate rapidly (Breiter et al. Reference Breiter, Etcoff, Whalen, Kennedy, Rauch, Buckner, Strauss, Hyman and Rosen1996; Büchel et al. Reference Büchel, Dolan, Armony and Friston1999; Fischer et al. Reference Fischer, Wright, Whalen, McInerney, Shin and Rauch2003; Whalen et al. Reference Whalen, Kagan, Cook, Davis, Kim, Polis, McLaren, Somerville, McLean, Maxwell and Johnstone2004; Wright et al. Reference Wright, Fischer, Whalen, McInerney, Shin and Rauch2001), suggesting that the amygdala is involved in attention to salient stimuli, but calling into question the idea that the amygdala is necessary to fear per se (for a similar point, see Adolphs Reference Adolphs2008; Reference Adolphs2010; Pessoa & Adolphs Reference Pessoa and Adolphs2010; Todd & Anderson Reference Todd and Anderson2009; Whalen Reference Whalen2007).Footnote 5

The amygdala's role in detecting motivationally salient stimuli would also explain why increased amygdala activity is observed in instances that do not involve the experience of fear, such as when stimuli are experienced as subjectively arousing (e.g., Bradley et al. Reference Bradley, Codispoti, Cuthbert and Lang2001; Weierich et al. Reference Weierich, Wright, Negreira, Dickerson and Barrett2010), intense (e.g., Bach et al. Reference Bach, Schachinger, Neuhoff, Esposito, Di Salle, Lehmann, Herdener, Scheffler and Seifrit2008), emotionally “impactful” (e.g., Ewbank et al. Reference Ewbank, Barnard, Croucher, Ramponi and Calder2009), or valuable (Jenison et al. Reference Jenison, Rangel, Oya, Kawasaki and Howard2011). Moreover, not all instances of fear are accompanied by increased amygdala activity (for a review, see Suvak & Barrett Reference Suvak and Barrett2011). For example, some behaviors that rats perform in dangerous contexts are not amygdala-dependent (e.g., avoiding the location of a threat: Vazdarjanova & McGaugh Reference Vazdarjanova and McGaugh1998; “defensive treading,” where bedding is kicked in the direction of the threat: Kopchia et al. Reference Kopchia, Altman and Commissaris1992). In humans, threatening contexts devoid of salient visual stimuli (e.g., preparing to give a speech in front of an audience), actually produce deactivatations in the amygdala (Wager et al. Reference Wager, van Ast, Hughes, Davidson, Lindquist and Ochsner2009a; Reference Wager, Waugh, Lindquist, Noll, Fredrickson and Taylor2009b). Moreover, electrical stimulation to the amygdala produces a range of experiences in humans, calling into question the idea that the amygdala is specifically linked to instances of fear (Bancaud et al. Reference Bancaud, Brunet-Bourgin, Chauvel and Halgren1994; Gloor Reference Gloor1990; Halgren et al. Reference Halgren, Walter, Cherlow and Crandall1978).

Our meta-analytic findings were inconsistent with a locationist hypothesis of amygdala function but were more consistent with the psychological constructionist hypothesis. Our density analyses revealed that, as compared to other brain regions, voxels within both amygdalae had more consistent increases in activation during instances of fear perception than during the perception of any other emotion category (Table 1). These voxels were not functionally specific for instances of perceiving fear, however. An insignificant χFootnote 2 analysis revealed that the voxels with consistent increases in activation during instances of fear perception were equally likely to have increased activity during instances of other emotion categories (see Fig. 6 for the proportion of study contrasts in the database for each emotion category that are associated with increased activity in right [R] amygdala; see Fig. S2 in the supplementary material, available at http://www.journals.cambridge.org/bbs2012008, for left [L] amygdala). Furthermore, instances of fear experience did not show a consistent increase in activation in either amygdala when compared to what would be expected by chance in other regions of the brain. Yet, as compared to other brain regions, voxels within bilateral amygdala had more consistent increases in activation during instances of disgust experience than during the experience of other emotion categories (Table 1). A χFootnote 2 analysis revealed that these voxels were functionally selective for of the experience of disgust, as there was more likely to be increased activity in those voxels during instances of disgust experience than during the experience of anger, fear, happiness or sadness (Table 2). Those voxels were not functionally specific to instances of disgust experience, however (Fig. 6; Fig. S2). Finally, as compared to other brain regions, a voxel in L. amygdala had more consistent increases in activation during instances of sadness perception than during the perception of other emotion categories (Table 1). An insignificant χFootnote 2 analysis revealed that this voxel was not functionally selective for instances of sadness, however.

Figure 6. Proportion of Study Contrasts with Increased Activation in Four Key Brain Areas. The y-axes plot the proportion of study contrasts in our database that had increased activation within 10mm of that brain area. The x-axes denote the contrast type separated by experience (exp) and perception (per). All brain regions depicted are in the right hemisphere. See Figures S2 and S3 in supplementary materials, available at http://www.journals.cambridge.org/bbs2012008, for additional regions. A color version of this image can be viewed in the online version of this target article at http://www.journals.cambridge.org/bbs.

Table 1. Brain Regions with a Consistent Increase in Activity Associated with the Experience or Perception of Discrete Emotion Categories in Density Analyses

Table 2. Brain Regions with a Consistent Increase in Activity Associated with the Experience or Perception of Discrete Emotion Categories in χFootnote 2 Analyses

Our logistic regressions confirmed and expanded upon our density and χFootnote 2 findings. There was more likely to be increased activity in the L. amygdala when participants were perceiving instances of fear or experiencing instances of disgust than when perceiving or experiencing any other emotion categories (Fig. 5; Table S6). These findings are consistent with the psychological constructionist hypothesis that the amygdala responds to salient perceptual stimuli because contrasts in our database that assessed the perception of fear and experience of disgust tended to use visual stimuli that are novel or unfamiliar to participants.Footnote 6 Findings for the R. amygdala also supported a psychological constructionist view. Increases in activity in the R. amygdala were likely when participants were experiencing or perceiving instances of any highly arousing emotion category (i.e., anger, disgust, fear) (Fig. 5; Table S6). There was likely to be no increase in activity in the L. amygdala when participants were focusing on their internal state (i.e., when emotion experience was induced via recall of a personal event and mental imagery; Fig. 5; Table S6). This finding replicates prior meta-analytic evidence (Costafreda et al. Reference Costafreda, Brammer, David and Fu2008) and is consistent with our hypothesis that the amygdala responds preferentially to salient exteroceptive (vs. interoceptive) sensations.

5.2. The anterior insula

Locationist accounts hypothesize that the anterior insula (Fig. 1, green) is the brain basis of disgust (e.g., Jabbi et al. Reference Jabbi, Bastiaansen and Keysers2008; Wicker et al. Reference Wicker, Keysers, Plailly, Royet, Gallese and Rizzolati2003; for reviews, see Calder et al. Reference Calder, Lawrence and Young2001; Calder Reference Calder2003) based on the belief that disgust evolved from a primitive food rejection reflex (Rozin et al. Reference Rozin, Haidt, McCauley, Lewis and Havilland-Jones2000) or bodily aversion to disease-threat (e.g., Curtis et al. Reference Curtis, Aunger and Rabie2004). Individuals with damage to the anterior insula and basal ganglia have difficulty perceiving instances of disgust in facial and vocal caricatures (Adolphs et al. Reference Adolphs, Tranel and Damasio2003; Calder et al. Reference Calder, Keane, Manes, Antoun and Young2000). They also report experiencing less disgust in response to scenarios about body products, envelope violation, and animals that typically evoke disgust in people with intact insulas (Calder et al. Reference Calder, Keane, Manes, Antoun and Young2000). Individuals with neurodegenerative diseases affecting the insula and basal ganglia (such as Huntington's and Parkinson's disease) also show diminished experiences of disgust to foul smelling odors (Mitchell et al. Reference Mitchell, Heims, Neville and Rickards2005) and have difficulty perceiving instances of disgust in the faces of others (e.g., Kipps et al. Reference Kipps, Duggins, McCusker and Calder2007; Sprenglemeyer et al. 1996; 1998; Suzuki et al. Reference Suzuki, Hoshino, Shigemasu and Kawamura2006; see Calder et al. Reference Calder, Lawrence and Young2001; Sprengelmeyer Reference Sprengelmeyer2007, for reviews), although the specificity of these findings remains in question (e.g., Calder et al. Reference Calder, Keane, Young, Lawrence, Mason and Barker2010; Milders et al. Reference Milders, Crawford, Lamb and Simpson2003). Patients who received electrical stimulation to the anterior insula reported visceral sensations consistent with (but not specific to) the experience of disgust (e.g., sensations in the stomach or throat, smelling or tasting something bad, nausea; Penfield & Faulk 1955).

In a psychological constructionist hypothesis, the anterior insula plays a key role in representing core affective feelings in awareness (Fig. 2, panel D, dark pink). The anterior insula is thought to be involved in the awareness of bodily sensations (Craig Reference Craig2002) and affective feelings (Craig Reference Craig2009). Sometimes sensations from the body are experienced as physical symptoms, but more often they are experienced as states that have some hedonic tone and level of arousal. Sometimes those affective feelings are experienced as emotion. To the extent that brain states corresponding to instances of disgust represent a stimulus's consequence for the body, then the anterior insula will show increased activation. Indeed, a key ingredient in the mental states labeled “disgust is likely a representation of how an object will affect the viscera. In support of a psychological constructionist view, anterior insula activation is observed in a number of tasks that involve awareness of body states, but not disgust per se. The anterior insula shows increased activation during awareness of body movement (e.g., Tsakiris et al. Reference Tsakiris, Hesse, Boy, Haggard and Fink2007), gastric distention (e.g., Wang et al. Reference Wang, Tomasi, Backus, Wang, Telang, Geliebter, Korner, Bauman, Fowler, Thanos and Volknow2008), and orgasm (e.g., Ortigue et al. Reference Ortigue, Grafton and Bianchi-Demicheli2007). Electrical stimulation of the insula produces sensations consistent with the category disgust, but it also produces a range of other visceral sensations including feelings of movement, twitching, warmth and tingling in the lips, tongue, teeth, arms, hands, and fingers (Penfield & Falk Reference Penfield and Falk1955). Dorsal anterior insula is also a hub in a large-scale network involved in what has been called a ventral attention system (Corbetta & Shulman Reference Corbetta and Shulman2002, Corbetta et al. Reference Corbetta, Patel and Shulman2008) that guides attention allocation and orienting (e.g., Eckert et al. Reference Eckert, Menon, Walczak, Ahlstrom, Denslow, Horowitz and Dubno2009). These findings again point to the idea that body-based sensory signals constitute a source of attention in the brain.

Our meta-analytic findings were inconsistent with the locationist account that the anterior insula is the brain seat of disgust but were more consistent with the psychological constructionist account that insula activity is correlated with interoception and the awareness of affective feelings. Our density analyses revealed that as compared to other brain regions, voxels within the right [R.] anterior insula had more consistent increases in activation during instances of disgust perception than during the perception of any other emotion category (Table 1). Our χFootnote 2 analysis revealed that only four of the voxels identified in the density analysis showed functional selectivity for instances of disgust perception (Table 2), however, and increased activity in R. insula was not specific to instances of disgust perception (Fig. 6). Our logistic regression findings for the R. anterior insula were consistent with the psychological constructionist hypothesis that the insula supports representation of core affective feelings. Increased activity in R. anterior insula was likely when participants were explicitly evaluating their feelings and representing them in awareness (Fig. 5, Table S6). Instances of disgust perception might consistently involve increased activation in the insula because people are more likely to simulate visceral states (such as those associated with the gut and food rejection) when perceiving facial behaviors characterized by a wrinkled nose and curled lip (i.e., oral revulsion; Angyal Reference Angyal1941; see also, Rozin et al. Reference Rozin, Haidt, McCaule, Lewis, Haviland-Jones and Barrett2008; von dem Hagen et al. Reference von dem Hagen, Beaver, Ewbank, Keane, Passamonti, Lawrence and Calder2009).

As compared to other brain regions, a greater spatial extent of voxels within the left [L.] anterior insula had consistent increases in activation during instances of disgust experience than during the experience of any other emotion category (Table 1). As compared to other brain regions, two voxels in L. anterior insula also had more consistent increases in activation during instances of anger experience than during the experience of any other emotion (Table 1; only one voxel showed functional selectivity, see Table 2). Our logistic regressions replicated this general finding. Increased activity in L. anterior insula was more likely when participants were experiencing an instance of anger than when they were experiencing any other emotion category (Fig. 5; Table S6). These findings, along with subsequent findings (see sect. 5.3 for the orbitofrontal cortex, sect. 5.6 for the anterior temporal lobe and ventrolateral prefrontal cortex, and 5.7 for the dorsolateral prefrontal cortex) suggest that instances of anger are associated with increased activity in a broad set of areas in the left frontal and temporal lobes.

5.3. The orbitofrontal cortex

Locationist accounts link the orbitofrontal cortex (OFC) to anger (Fig. 1, rust), although the OFC is a large structure and has admittedly been linked to many other psychological phenomena. Primary support for the OFC-anger hypothesis derives from prior meta-analytic reviews of the neuroimaging literature (Murphy et al. Reference Murphy, Nimmo-Smith and Lawrence2003; Vytal & Hamann Reference Vytal and Hamann2010). Studies using electroencephalography (EEG) also associate instances of anger with the prefrontal cortex (PFC).Footnote 7 Using EEG, activity in the left PFC is associated with instances of anger experience in response to an insult (Harmon-Jones & Sigelman Reference Harmon-Jones and Sigelman2001) and with the personality disposition to experience angry feelings (Harmon-Jones & Allen Reference Harmon-Jones and Allen1998). Other evidence for an OFC-anger hypothesis is more circumstantial. For instance, there is a body of evidence linking the OFC to aggression. It is far from clear that aggression is an unambiguous index of the entire category of anger, however. Nonhuman animals aggress in a number of different contexts (e.g., maternal aggression, sexual aggression, predatory aggression, defensive aggression; Moyer Reference Moyer1968); only some of which are associated with the concept called “anger” in English. Humans do a number of things in anger, only some of which constitute aggression. With that caveat, there is certainly evidence linking increased activity in the OFC to aggression. EEG activity in the left PFC is associated with an increased tendency to retaliate towards another person following an insult (by allocating him or her a dose of unpleasant hot sauce in a putative taste test; Harmon-Jones & Sigelman Reference Harmon-Jones and Sigelman2001). Aggressive behavior in rats is associated with increased activity in the ventral forebrain (including the OFC) (Ferris et al. Reference Ferris, Stolberg, Kulkarni, Murugavel, Blanchard, Blanchard, Febo, Brevard and Simon2008). Some lesion evidence is consistent with the idea that the OFC produces aggression in monkeys, because OFC lesions reduce aggression (towards humans: Butter & Snyder Reference Butter and Snyder1972; Kamback Reference Kamback1973; towards other monkeys: Raleigh et al. Reference Raleigh, Steklis, Ervin, Kling and McGuire1979). The majority of lesion studies find that monkeys (e.g., Deets et al. Reference Deets, Harlow, Singh and Blomquist1970; Machado & Bachevalier Reference Machado and Bachevalier2006; Raleigh et al. Reference Raleigh, Steklis, Ervin, Kling and McGuire1979) and rats (de Bruin et al. Reference de Bruin, Van Oyen and Van De Poll1983) are more aggressive towards conspecifics following OFC lesions, however. Similarly, electrical stimulation of the lateral OFC (lOFC; in cats: Siegel et al. Reference Siegel, Edinger and Dotto1975) and the medial OFC (mOFC; in cats: Siegel et al. Reference Siegel, Edinger and Lowenthal1974; and in rats: de Bruin Reference de Bruin1990) inhibits, rather than causes, aggressive behavior. Humans with lesions in the ventromedial PFC (which contains the OFC) become frustrated more easily and engage in more verbal (but not physical) aggression than do neurologically intact subjects (Grafman et al. Reference Grafman, Schwab, Warden, Pridgen, Brown and Salazar1996). Psychopathy and antisocial disorder are marked by increased aggression and correspond to structural (e.g., Raine et al. Reference Raine, Lencz, Bihrle, LaCasse and Colletti2000) and functional (e.g., Glenn et al. Reference Glenn, Raine and Schug2009; Harenski et al. Reference Harenski, Kim and Hamann2009) changes to the mOFC (Yang & Raine Reference Yang and Raine2009). Fewer studies have linked the lOFC to aggressive behavior in humans, but one study found that individuals with borderline personality disorder who have lowered baseline lOFC (BA 47) activity are more likely to aggress against others (Goyer et al. Reference Goyer, Konicki, Schulz and Silk1994).

A psychological constructionist view hypothesizes that portions of the OFC play a role in core affect as a site that integrates exteroceptive and interoceptive sensory information to guide behavior. Together, sensory information from the world and sensory information from the body guide an organism's response to the environment and allow it to engage in behavior that is well tuned to the context (defined both by the external surroundings and the organism's goals). With the lOFC's connections to sensory modalities (e.g., Barbas Reference Barbas1988; Rolls Reference Rolls1999, see Kringelbach & Rolls Reference Kringelbach and Rolls2004) and the mOFC's connections to areas involved in visceral control (e.g., Carmichael & Price Reference Carmichael and Price1995; Eblen & Graybiel Reference Eblen and Graybiel1995; Ongür & Price 1998; Reference Öngür and Price2000; Rempel-Clower & Barbas Reference Rempel-Clower and Barbas1998; see Kringelbach & Rolls [2004] for a review), the OFC is anatomically well suited to perform this role. We are not claiming that this is the OFC's specific function, but only that it is a brain region that is important to realizing this function. Consistent with the idea that the OFC unites internal and external sensory information, the lOFC and the mOFC have been linked to associative learning (Rolls et al. Reference Rolls, Hornak, Wade and McGrath1994; Reference Rolls, Critchley, Mason and Wakeman1996) decision making (e.g., Bechara et al. Reference Bechara, Tranel, Damasio and Damasio1996; Reference Bechara, Tranel and Damasio2000; Koenigs et al. Reference Koenigs, Young, Adolphs, Tranel, Cushman, Hauser and Damasio2007) and reversal learning, in which the reward values associated with choice options are reversed and animals must learn the current reward value (Chudasama & Robbins Reference Chudasama and Robbins2003; Hornak et al. Reference Hornak, O'Doherty, Bramham, Rolls, Morris, Bullock and Polkey2004; Rudebeck & Murray Reference Rudebeck and Murray2008). The inability to properly integrate exteroceptive and interoceptive information will result in behavior that is inappropriate to that context, explaining the altered non-aggressive social behavior (e.g., Beer et al. Reference Beer, John, Scabini and Knight2003; Eslinger & Damasio Reference Eslinger and Damasio1985; Saver & Damasio Reference Saver and Damasio1991; see Damasio et al. Reference Damasio, Tranel and Damasio1990) and aggression (Grafman et al. Reference Grafman, Schwab, Warden, Pridgen, Brown and Salazar1996) observed in individuals with OFC damage.

Our meta-analytic findings were inconsistent with the locationist hypothesis that the OFC is the brain seat of anger. As compared to voxels within other brain regions, voxels within the OFC did not have more consistent increases during instances of anger experience or perception than during any other emotion category. Rather, as compared to voxels within other brain regions, voxels within the left lOFC had more consistent increases in activation during instances of disgust experience than during the experience of other emotion categories (Table 1). Voxels within the right lOFC, as compared to voxels within other brain regions, had more consistent increases in activation during instances of disgust perception than during the perception of other emotion categories (Table 1). χFootnote 2 analyses indicated that there was some functional selectivity for instances of disgust experience and perception in the voxels in the right and left lOFC that were respectively identified in the density analysis. Activity in the right and left lOFC was not specific to instances of disgust experience or perception, however (Fig. 6; Fig. S2). Our logistic regressions confirmed that when participants were perceiving an instance of disgust, there was more likely to be increased activity in the right lOFC than when participants were perceiving instances of any other emotion category (Fig. 5; Table S6).

Our logistic regressions revealed that increased activity in the left lOFC was more likely when participants were experiencing instances of anger than when experiencing instances of any other emotion category (Fig. 5; Table S6). Although in and of itself, this finding provides partial support for the OFC-anger hypothesis, our other meta-analytic findings indicate that increased activity in the left hemisphere during instances of anger is not restricted to the OFC, or even the prefrontal cortex (see sect. 5.2, “The anterior insula,” sect. 5.6, “Anterior temporal lobe and ventrolateral prefrontal cortex,” and sect. 5.7, “Dorsolateral prefrontal cortex”). Additionally, our logistic regressions revealed that increased activity in the left and right lOFC was likely when participants were experiencing a range of exteroceptive (auditory stimuli, visual pictures) and interoceptive (experience or perception of high-arousal core affect) sensations (Fig. 5; Table S6). This finding is consistent with the psychological constructionist hypothesis that OFC plays a more general role in integrating heteromodal sensations.

5.4. The anterior cingulate cortex

Locationist accounts hypothesize that pregenual anterior cingulate cortex (pACC; BAs 24, 32) and subgenual anterior cingulate cortex (sACC; BA 25) are the brain basis of sadness (Fig. 1, blue). The pACC and sACC have known affective function and are thought to instantiate the visceromotor responses observed during classical conditioning, pain, and affective behaviors (Devinksy et al. 1995; Vogt Reference Vogt2005). The ACC-sadness hypothesis, like the OFC-anger hypothesis, derives support from prior meta-analyses of the neuroimaging literature (e.g., Murphy et al. Reference Murphy, Nimmo-Smith and Lawrence2003; Phan et al. Reference Phan, Wager, Taylor and Liberzon2002). In the behavioral neuroscience literature, pACC has been linked to sadness because of its role in producing the vocalizations that infant animals make when separated from their mother (e.g., MacLean & Newman Reference MacLean and Newman1988; see Panksepp Reference Panksepp1998; Reference Panksepp2007); the link between infant vocalizations and unpleasant affect are in question, however (Blumberg & Sokoloff Reference Blumberg and Sokoloff2001). One study in humans found that pACC lesions (including lesions to dorsomedial prefrontal cortex) produce hypersensitivity and an increased tendency to cry at sad events (Hornak et al. Reference Hornak, Bramham, Rolls, Morris, O'Doherty, Bullock and Polkey2003). If the pACC were involved in instantiating instances of sadness, then lesions to this structure should abolish the tendency to cry at sad events. These findings are therefore more consistent with the idea that pACC regulates instances of sadness. Finally, pACC is implicated in pain, perhaps because it supports the feeling of “suffering” (cf. Vogt Reference Vogt2005). The sACC, on the other hand, has been linked to sadness because of its role in depression. Clinical depression is marked by structural and functional changes in sACC (see Gotlib & Hamilton [2008] for a review), although clinical depression involves many symptoms above and beyond the experience of sadness (Coyne Reference Coyne1994). Electrical stimulation of the sACC relieves intractable depression by reducing feelings of apathy and anhedonia, normalizing sleep disturbances, and decreasing gross motor impairments (e.g., Mayberg et al. Reference Mayberg, Lozano, Voon, McNeely, Seminowicz, Hamani, Schwalb and Kennedy2005).

Our psychological constructionist hypothesis draws on the neuroscience literature showing that pACC, sACC, and the more dorsal anterior midcingulate cortex (aMCC) (Vogt Reference Vogt, Vogt and Gabriel1993; Vogt et al. Reference Vogt, Berger and Derbyshire2003) take part in distinct psychological operations related to realizing core affect. In our view, the pACC and sACC (along with the adjacent posterior mOFC) are cortical sites for visceral regulation that help to realize a core affective state during motivated action (Devinksy et al. 1995) (Fig. 2, panel B, light pinks). We would therefore predict increased activation in pACC and sACC across a variety of emotional instances. The hypothesized role of sACC in regulating somatovisceral states could explain why sACC is involved in the affective changes that accompany both depression (e.g., Drevets et al. 1992; see Gotlib & Hamilton Reference Gotlib and Hamilton2008) and mania (e.g., Fountoulakis et al. Reference Fountoulakis, Giannakopoulos, Kövari and Bouras2008), and why electrical stimulation of this region helps to relieve chronic depression (Mayberg, et al. Reference Mayberg, Lozano, Voon, McNeely, Seminowicz, Hamani, Schwalb and Kennedy2005). The aMCC (Fig. 2, panel B, dusty pink) is hypothesized to play a role in executive attention and motor engagement during response selection (Mansouri et al. Reference Mansouri, Tanaka and Buckley2009; Rushworth et al. Reference Rushworth, Buckley, Behrens, Walton and Bannerman2007). In this view, aMCC delivers sources of exteroceptive sensory information (from thalamic projections; Barbas et al. Reference Barbas, Henion and Dermon1991) and internal sensory information (from the insula; Mesulam & Mufson Reference Mesulam and Mufson1982) to direct attention and motor responses (via projections to lateral PFC and the supplementary motor area; e.g., Barbas & Pandya Reference Barbas and Pandya1989; for reviews, see Devinsky et al. Reference Devinsky, Morrell and Vogt1995; Paus Reference Paus2001). These anatomical connections can explain why the aMCC is part of an intrinsic brain network that shows increased activity when stimuli in the environment are personally salient (Seeley et al. Reference Seeley, Menon, Schatzberg, Keller, Glover, Kenna, Reiss and Greicius2007). They also explain why the aMCC is responsible for resolving action selection during situations involving conflicting sensory information (e.g., Grinband et al. Reference Grinband, Savitsky, Wager, Teichert, Ferrera and Hirsch2011; Milham et al. Reference Milham, Banich, Webb, Barad, Cohen, Wszalek and Kramer2001; Nelson et al. Reference Nelson, Reuter-Lorenz, Sylvester, Jonides and Smith2003; see Banich Reference Banich2009; Shackman et al. Reference Shackman, Salomons, Slagter, Fox, Winter and Davidson2011; van Snellenberg & Wager 2009). By extension, as a visceromotor (i.e., autonomic) control area sACC/pACC might be involved in resolving which sensory inputs influence the body when there are multiple sources of sensory input that can influence an organism's body state.

Our meta-analytic evidence is inconsistent with the locationist account that the ACC is the brain basis of sadness, but more consistent with a psychological constructionist hypothesis of ACC function. As compared to voxels within other brain regions, voxels within the sACC, pACC and aMCC did not have more consistent increases when participants were experiencing or perceiving instances of sadness than during any other emotion category (Fig. 6). As compared to voxels within other brain regions, a greater spatial extent of voxels within the aMCC had consistent increases in activation during instances of fear perception than the perception of any other emotion category (Table 1). The amygdala, which responds to motivationally salient exteroceptive sensory stimuli (see sect. 5.1 “The amygdala”), projects to this area of aMCC (Vogt & Pandya 1987), so it is possible that increased activity here reflects response preparation to salient stimuli in the environment.

Our logistic regressions revealed that increases in sACC were likely when participants were engaged in cognitive load (Fig. 5; Table S6). Cognitive load typically occurred in studies where participants were asked to attend to core affective feelings (e.g., focus on their feelings, rate their feelings) or affective stimuli (e.g., focus on an affective stimulus, rate a stimulus' emotional value) and so it is possible that this finding is indicative of the sACC's role in instantiating core affective feelings. Increased activity in the sACC was marginally (p < 0.09) likely when participants were evaluating their feelings, again consistent with this area's role as a visceromotor regulation site. Finally, consistent with a response selection hypothesis of aMCC function, increased activity in the aMCC was likely when participants were engaged in cognitive load (Fig. 5; Table S6).

5.5. Dorsomedial prefrontal cortex, medial temporal lobe, and retrosplenial cortex/posterior cingulate cortex

Our psychological constructionist approach hypothesizes that a range of other brain regions are important to realizing instances of emotion experience and perception, including dorsomedial prefrontal cortex (DMPFC), ventromedial prefrontal cortex (VMPFC), medial temporal lobe (MTL), and retrosplenial cortex/posterior cingulate cortex (PCC) (Fig. 2, panel B, purples), which are associated with the psychological operation of conceptualization. As part of the process of making meaning out of sensory cues, we hypothesize that these brain areas use stored representations of prior experiences to make meaning of core affective inputs that come from the self or observing others. Locationist views do not hypothesize specific roles for these brain regions in emotion because they are usually considered to have a “cognitive” function, insofar that they support memory (Vincent et al. Reference Vincent, Snyder, Fox, Shannon, Andrews, Raichle and Buckner2006), object perception (Bar 2009), and theory of mind (Mitchell Reference Mitchell2009). In our view, these brain regions should not necessarily be more involved in instances of one category of emotion than another, although we would expect them to be part of the more general neural reference space for discrete emotion.

As we predicted, our meta-analytic results revealed that regions of the conceptualization network such as DMPFC, MTL, and retrosplenial cortex (Buckner et al. Reference Buckner, Andrews-Hanna and Schacter2008) were part of the neural reference space for discrete emotion (Fig. 4).Footnote 8 Our findings indicate that the conceptualization network is integral in the experience and perception of discrete emotions, and are consistent with the hypothesis (in Barrett Reference Barrett2006b; Reference Barrett2009a; Reference Barrett2009b) that prior episodic experience helps shape experienced or perceived affect into meaningful instances of emotion.

Despite general involvement in emotion (Fig. 1; Fig. S3), there was some functional selectivity for instances of certain emotion categories in hubs within the conceptualization network. For instance, across our density analyses and logistic regressions, we found that instances of the experience of sadness (Tables 1 & 2) and experience of happiness (Table 1) were each associated with relatively greater consistent increases in activation in areas of DMPFC than other emotion categories. Consistent with the role of conceptualization in simulating episodic experience (Schacter et al. Reference Schacter, Addis and Buckner2007), our logistic regressions revealed that increased activity in DMPFC was likely when participants were engaging in emotion inductions involving recall and films (Fig. 5). Increased activity in some clusters of DMPFC was likely when participants were perceiving emotion in faces, bodies or voices. This finding is consistent with the psychological constructionist hypothesis that the conceptualization network is brought to bear when affective facial behaviors are perceived as emotional. Other clusters of DMPFC showed an opposite pattern: When participants were perceiving emotion, increased activity in DMPFC was not likely (Table S6). Just as perception of others and self-referential thinking involve overlapping yet distinct aspects of DMPFC (Ochsner et al. Reference Ochsner, Knierim, Ludlow, Hanelin, Ramachandran and Mackey2004a), some aspects of DMPFC might be functionally selective for conceptualization during emotion perception whereas others support conceptualization during emotion experience.

Several emotion categories were also associated with consistent increases in activation in the MTL (Tables 1 & 2; see Table S6). Our logistic regressions revealed that, as in the DMPFC, increased activity in the right hippocampus was likely to occur when participants perceived any instance of emotion in a face, body, or voice. Increased activity in the left hippocampus, on the other hand, was likely to occur when participants perceived instances of fear (Table S6). This finding is more likely to be related to the encoding of salient stimuli in memory than simulating prior experiences, as the amygdala also had increased activity during instances of fear perception and is known to have functional connectivity with the hippocampus during encoding of salient stimuli (Kensinger & Corkin Reference Kensinger and Corkin2004).

5.6. Anterior temporal lobe and ventrolateral prefrontal cortex

According to a psychological constructionist account, networks supporting language (e.g., Vigneau et al. Reference Vigneau, Beaucousin, Hervé, Duffau, Crivello, Houdé, Mazoyer and Tzourio-Mazoyer2006) should consistently show increased activity during instances of emotion experience and perception as linguistically-grounded concepts are brought to bear to make meaning of core affective feelings. In locationist accounts, language is thought to be epiphenomenal to discrete emotion (Ekman & Cordaro Reference Ekman and Cordaro2011), although recent behavioral studies show that categorical perception of discrete emotion is supported by language (Fugate et al. Reference Fugate, Gouzoules and Barrett2010; Roberson & Davidoff Reference Roberson and Davidoff2000; Roberson et al. Reference Roberson, Damjanovic and Pilling2007).

Consistent with the psychological constructionist view, nodes within networks supporting language were part of the neural reference space for discrete emotion (Fig. 4) In particular, the anterior temporal lobe (ATL) and ventrolateral prefrontal cortex (VLPFC) (Fig. 2, Panels A, B, C, green) had consistent increases in activity across studies of discrete emotion. The ATL supports language as a heteromodal association area involved in the representation of concepts (Lambon Ralph et al. Reference Lambon Ralph, Pobric and Jefferies2009; Pobric et al. Reference Pobric, Jefferies and Ralph2007; Rogers et al. Reference Rogers, Lambon Ralph, Garrard, Bozeat, McClelland, Hodges and Patterson2004) and the right ATL has been implicated in the representation of abstract social concepts (e.g., Zahn et al. Reference Zahn, Moll, Paiva, Garrido, Krueger, Huey and Grafman2009). Patients with semantic dementia have focal atrophy to the ATL, difficulty utilizing semantic knowledge, and exhibit deficits in emotion perception (Rosen et al. Reference Rosen, Pace-Savitsky, Perry, Kramer, Miller and Levenson2004) and empathy (Rankin et al. Reference Rankin, Gorno-Tempini, Allison, Stanley, Glenn, Weiner and Miller2006). Areas of the VLPFC, on the other hand, are implicated in semantic processing tasks (e.g., Gitelman et al. Reference Gitelman, Nobre, Sonty, Parrish and Mesulam2005), categorization of objects (e.g., Freedman et al. Reference Freedman, Riesenhuber, Poggio and Miller2001), representation of feature-based information for abstract categories (e.g., Freedman et al. Reference Freedman, Riesenhuber, Poggio and Miller2002; see Miller et al. Reference Miller, Freedman and Wallis2002), selection amongst competing response representations (e.g., Badre & Wagner Reference Badre and Wagner2007; Schnur et al. Reference Schnur, Schwartz, Kimberg, Hirshorn, Coslett and Thompson-Schill2009), and inhibition of responses (Aaron et al. Reference Aron, Robbins and Poldrack2004). It is therefore not clear that the VLPFC's role is functionally specific to language, but it is certainly functionally selective for language in certain instances. The VLPFC also helps comprise the ventral frontoparietal network that is thought to be involved in directing attention to salient stimuli in the environment (Corbetta & Shulman Reference Corbetta and Shulman2002; Corbetta et al. Reference Corbetta, Patel and Shulman2008), suggesting a more general role for this region in executive attention.

As compared to other brain regions, voxels within the ATL did not have more consistent increases during instances of one emotion category than others (Fig. S3). Our logistic regressions suggested that increased activity in the left ATL was more likely when participants were experiencing an instance of anger than any other emotion category, however (Fig. 5; Table S6). Instances of anger experience thus involve areas throughout the left frontal and temporal lobes (see sect. 5.2, “The anterior insula,” sect. 5.3, “The orbitofrontal cortex,” and sect. 5.7, “Dorsolateral prefrontal cortex”). Increased activity in the right ATL, on the other hand, was likely when participants were evaluating a stimulus (i.e., determining the emotional meaning of a face, voice, or picture; Table S6). This finding is consistent with the hypothesis that language is brought to bear when constructing emotional percepts from exteroceptive sensations.

Our density analyses revealed that as compared to voxels within other brain regions, voxels within the right VLPFC had more consistent increases during instances of disgust perception than during the perception of any other emotion category; these findings were confirmed with a logistic regression (Table 1; Table S6). χFootnote 2 analyses revealed that a subset of the voxels identified in the density analysis were functionally selective for instances of disgust perception (Table 2), although they were not specific to instances of disgust (Fig. S3). The most common finding across our logistic regressions linked increases in the left VLPFC to instances in which participants were explicitly paying attention to emotional information (Fig. 5; Table S6). As in other left frontal and temporal areas (see sects. 5.2, “The anterior insula,” 5.3, “The orbitofrontal cortex,” 5.6, “Anterior temporal lobe and ventrolateral prefrontal cortex”), increased activity in the left VLPFC was likely when participants were experiencing or perceiving instances of anger.

5.7. Dorsolateral prefrontal cortex

According to a psychological constructionist account, networks supporting executive attention (see Miller & Cohen Reference Miller and Cohen2001; Petrides Reference Petrides2005; for a meta-analysis, see Wager & Smith Reference Wager and Smith2003) should consistently show increased activity during instances of emotion experience and perception because executive attention directs other psychological operations during the construction of emotion. Locationist accounts do not propose specific roles for these networks in emotion, although they might allow that networks supporting executive attention take part in regulation of emotion after it is generated (as in Ochsner et al. Reference Ochsner, Ray, Cooper, Robertson, Chopra, Gabrieli and Gross2004b; Urry et al. Reference Urry, van Reekum, Johnstone, Kalin, Thurow, Schaefer, Jackson, Frye, Greischar, Alexander and Davidson2006).Footnote 9

Consistent with our psychological constructionist hypothesis, nodes within networks supporting executive attention were part of the neural reference space for discrete emotion. In particular, the VLPFC and dorsolateral prefrontal cortex (DLPFC) (Fig. 2, Panel A, orange and striped green/orange) had consistent increases in activity across studies of discrete emotion. The DLPFC is part of the dorsal frontoparietal network that is thought to be involved in top-down, goal-directed selection for responses (Corbetta & Shulman Reference Corbetta and Shulman2002; Corbetta et al. Reference Corbetta, Patel and Shulman2008). Consistent with this functional connectivity, bilateral DLPFC is known to be involved in working memory (e.g., Champod & Petrides Reference Champod and Petrides2007; Constantinidis et al. Reference Constantinidis, Williams and Goldman-Rakic2002) and in the goal-directed control of attention (e.g., Rainer et al. 1998; see Miller Reference Miller2000).

We predicted that DLPFC would be part of the neural reference space for emotion (Fig. 4) because these voxels would be active during mental states in which participants attended to emotional feelings or perceptions (i.e., when participants had to hold affective information in mind in order to categorize it). Consistent with this prediction, increased activity in the right DLPFC was likely when participants were explicitly evaluating stimuli (Fig. 5; Table S6). Our density analyses also revealed that as compared to voxels within other brain regions, voxels in the right DLPFC were more likely to have increased activity during instances of anger perception than any other emotion categories (Table 1). Our χFootnote 2 analyses indicated that some of these voxels were functionally selective to perceiving instances of anger (Table 2), although they were not specific to perceiving instances of anger (Fig. S3).

5.8. The periaqueducal gray

The periaqueducal gray (PAG) is involved in regulating the autonomic substrates that allow for behavioral adaptations such as freezing, fleeing, vocalization, and reproductive behavior (e.g., Carrive et al. Reference Carrive, Bandler and Dampney1989; Behbehani Reference Behbehani1995; Gregg & Siegel Reference Gregg and Siegel2001; Kim et al. Reference Kim, Rison and Fanselow1993; Lovick Reference Lovick1992; Mobbs et al. Reference Mobbs, Petrovic, Marchant, Hassabis, Weiskopf, Seymour, Dolan and Frith2007; Van der Horst & Holstege Reference Van der Horst and Holstege1998) and also sends projections back to cortical sites involved in the regulation of visceral activations in the body (An et al. Reference An, Bandler, Ongur and Price1998; Mantyh Reference Mantyh1983). It is believed that certain adaptations are associated with certain emotion categories (e.g., animals freeze in fear, aggress in anger) but such links are far from empirically clear. Humans (like other mammals) do many things during instances of anger, for example. Sometimes humans yell, sometimes they hit, sometimes they remain very still, and sometimes they smile. Even rats do many things within a single emotion category: In the face of a threat, rats can freeze (e.g., LeDoux et al. Reference LeDoux, Ciccetti, Xagoraris and Romanski1990), flee (Vazdarjanova & McGaugh Reference Vazdarjanova and McGaugh1998), or engage in “defensive treading,” where they kick bedding in the direction of a known threat (Reynolds & Berridge Reference Reynolds and Berridge2002; Reference Reynolds and Berridge2003; Reference Reynolds and Berridge2008). In all these instances, PAG activity and the associated autonomic states it produces, are yoked to the action, and not to the emotion category. As a result, a psychological constructionist approach views PAG activity as nonspecifically involved in instances of emotion. A locationist approach has linked the PAG to distinct circuits corresponding to several emotion categories: rage, fear, joy, distress, love and lust (Panksepp Reference Panksepp1998). In a psychological construction approach, the assumption is that a given dedicated circuit for a specific behavioral adaptation (e.g., withdrawal) will be active across a range of emotion categories (e.g., a person can withdraw in instances of both fear and anger), and different dedicated circuits within the PAG (e.g., fight, flight) will be active within instances of a single emotion category depending upon which behavioral adaptation is more relevant for the immediate context.

Testing any hypothesis about the specificity of a subcortical region like the PAG is practically impossible, given the spatial and temporal limitations of brain imaging. Still, it is instructive to note that the PAG was consistently activated within the neural reference space for discrete emotion (even though subjects were lying still and not engaging in any overt physical action; see Fig. S4 in the supplementary materials, available at http://www.journals.cambridge.org/bbs2012008). Moreover, we did not find evidence of functional specificity for the PAG in our meta-analysis. It is possible that given the resolution problems, different circuits within the PAG were specifically active for discrete emotions. That said, it is also entirely plausible from the behavioral data that humans, like animals, perform a range of actions within a single category, and perform the same action at times across categories (even if it does not match our stereotypes of emotion–action links). In this meta-analysis, increased activity in the PAG did not correspond to any particular emotion category more than another (although instances of the experience of sadness were associated with consistent increases in activation in one voxel within ventral PAG; Tables 1 & 2). The logistic regressions demonstrated that increased activity in the PAG was likely when participants were experiencing or perceiving an instance of any high-arousal emotion category (Table S6). Consistent with this finding, activity in a separate cluster of PAG was likely when participants were experiencing instances of fear (Table S6). In a previous meta-analysis, we found that the PAG was most likely to have increased activity during unpleasant emotions (Wager et al. Reference Wager, Barrett, Bliss-Moreau, Lindquist, Duncan, Kober, Joseph, Davidson and Mize2008). Since all high-arousal emotion categories in our database were unpleasant (e.g., fear, anger, disgust), our present findings are consistent with Wager et al. (Reference Wager, Barrett, Bliss-Moreau, Lindquist, Duncan, Kober, Joseph, Davidson and Mize2008). Future research should probe whether the PAG preferentially shows increased activity during unpleasant states, highly aroused states, or during states that are both unpleasant and highly aroused.

5.9. Visual cortex

From our psychological construction approach, we would not be surprised to observe that voxels within visual cortex (or any sensory modality) show increased activation during emotion. In its most basic form, our hypothesis is that emotion emerges as a situated conceptualization of internal sensations from the body and external sensations from the world to create a unified conscious experience of the self in context. In fact, regions of visual cortex were some of the most frequent to appear in our meta-analytic findings (replicating several recent meta-analyses; e.g., Fusar-Poli et al. Reference Fusar-Poli, Placentino, Carletti, Landi, Allen, Surguladze, Benedetti, Abbamonte, Gasparotti, Barale, Perez, McGuire and Politi2009; Kober et al. Reference Kober, Barrett, Joseph, Bliss-Moreau, Lindquist and Wager2008; Vytal & Hamann Reference Vytal and Hamann2010). It is beyond the scope of this article to discuss these findings in detail, but we found that instances of fear experience, anger perception, and disgust experience were consistently associated with increased activity in regions of visual cortex ranging from V2 to visual association cortex (Tables 1 & 2). Our logistic regressions revealed that activation in visual cortex was not merely a by-product of the methods used. Although increased activity in visual cortex was likely when visual methods were used (e.g., pictures, faces, studies of perception; see Table S6), it was also likely in studies of unpleasant, high-arousal emotions (e.g., fear, disgust, unpleasant emotions more generally; Table S6). Together, these findings indicate that experiences or perceptions of unpleasant emotion categories are realized by brain states that include increased activity in visual cortex. See Gendron et al. (in preparation) for a discussion.

6. Conclusion

Over a century ago, William James wrote, “A science of the relations of mind and brain must show how the elementary ingredients of the former correspond to the elementary functions of the latter,” (James Reference James1890/1998, p. 28). James believed that emotions, thoughts, and memories are categories derived from commonsense with instances that do not require special brain centers. With respect to emotion, he wrote, “sensational, associational, and motor elements are all that [the brain] need contain” to produce the variety of mental states that correspond to our commonsense categories for emotion (cf. James 1890/1998, p. 473). James' view foreshadowed modern psychological constructionist models of the mind and the findings of our meta-analytic review, which are largely in agreement with this approach. Our findings are consistent with the idea that emotion categories are not natural kinds that are respected by the brain. The fact that some of the regions we report also appear in meta-analyses of other task domains (e.g., action simulation and perception, Grezes & Decety Reference Grezes and Decety2001; autobiographical memory, Svoboda et al. Reference Svoboda, McKinnon and Levine2006; decision making, Krain et al. Reference Krain, Wilson, Arbuckle, Castellanos and Milham2006; executive control, Owen et al. Reference Owen, McMillan, Laird and Bullmore2005; Wager & Smith Reference Wager and Smith2003; Wager et al. Reference Wager, Reading and Jonides2004; language, Vigneau et al. Reference Vigneau, Beaucousin, Hervé, Duffau, Crivello, Houdé, Mazoyer and Tzourio-Mazoyer2006; self-referential processing, Northoff 2006) means that these regions are not specific to emotion per se, and are also involved in constituting other cognitive and perceptual events (for a discussion of domain general networks, see Dosenbach et al. Reference Dosenbach, Visscher, Palmer, Miezin, Wenger, Kang, Burgund, Grimes, Schlaggar and Petersen2006; Nelson et al. Reference Nelson, Dosenbach, Cohen, Wheeler, Schlaggar and Petersen2010; Spreng et al. Reference Spreng, Mar and Kim2009; van Snellenberg & Wager 2009). Such findings show that even categories like emotion, cognition, and perception are not respected by the brain (Barrett Reference Barrett2009a; Duncan & Barrett Reference Duncan and Barrett2007; Pessoa Reference Pessoa2008).

In keeping with James' predictions, our meta-analytic review did not find strong evidence for a locationist hypothesis of brain–emotion correspondence (see Table 3 for a summary of findings). In all instances where a brain region showed consistent increases in activation during instances of a discrete emotion category (e.g., the amygdala in instances of fear perception), this increase was not specific to that category, failing to support a key locationist assumption. Some brain regions showed functional selectivity for instances of certain emotion categories; these findings perhaps point to differences in the contents of mental states (e.g., instances of anger experience often involve approach motivation, instances of disgust perception often involve simulation of bodily activation, and instances of fear perception often involve detection of unusual and hence salient stimuli).

Table 3. Summary of Brain Regions Showing Consistent Increases in Activation During Mental States and Methodological Manipulations

Mental states include states related to experiencing vs. perceiving discrete emotions, the experience and perception of affect, the experience or perception of individual discrete emotion categories, and mental states related to method types, stimulus types and other psychological variables. Brain regions consistently associated with mental states in the χ 2 analyses and logistic regressions are listed. Mental state-brain associations observed in the χ 2 analyses are printed in regular font. Logistic regression findings are in bold face font. Mental state-brain region associations observed in both types of analyses are marked with an asterisk (*).

Our meta-analytic findings were relatively more consistent with the psychological operations that we have considered ingredients of emotion here and in other papers (e.g., Barrett Reference Barrett2006b; 2009; Barrett et al. Reference Barrett, Lindquist, Bliss-Moreau, Duncan, Gendron, Mize and Brennan2007a; Kober et al. Reference Kober, Barrett, Joseph, Bliss-Moreau, Lindquist and Wager2008; Lindquist & Barrett Reference Lindquist and Barrett2008a; Wager et al. Reference Wager, Barrett, Bliss-Moreau, Lindquist, Duncan, Kober, Joseph, Davidson and Mize2008). In Kober et al. (Reference Kober, Barrett, Joseph, Bliss-Moreau, Lindquist and Wager2008), we hinted at the existence of basic psychological operations in the psychological construction of emotion. In other theoretical discussions (Barrett 2009) we explicitly hypothesized the need for mid-level scientific categories that describe the most basic psychological ingredients of the mind by referencing both biology and folk psychology when explaining how mental states like emotion experiences and perceptions arise (for a similar view, see Cacioppo et al. Reference Cacioppo, Berntson and Nusbaum2008). This target article is the first to investigate the extent to which brain regions associated with basic psychological domains show consistent increases in activation in neuroimaging studies of discrete emotion categories, despite a range of methodological variables. Of course, more work needs to be done to hone and refine our conceptions of the operations that are most psychologically primitive and to map them to networks in the brain, but this is a start.

Most notably, we observed consistent increases in activation in the brain regions implicated in conceptualization (simulation of prior episodic experiences), language (representation and retrieval of semantic concepts), and executive attention (volitional attention and working memory), suggesting that these more “cognitive” functions play a routine role in constructing experiences and perceptions of emotion. For example, increased activation in the DMPFC was observed when participants perceived instances of emotion on others' faces. Increased activation in the ATL was observed when participants focused on emotional stimuli. Increased activity in the VLPFC occurred when participants focused on the affective content of feelings or perceived instances of emotion on another person's face. Increased activity in the DLPFC occurred when participants evaluated the emotional content of a stimulus. One interpretation of these findings is that they are merely the result of the types of psychological tasks participants are asked to perform in the scanner during neuroimaging studies of emotion (e.g., recall, labeling, response selection), and that because of the limits of neuroimaging, these influences cannot be separated from an emotion itself. Yet, all data in our meta-analysis were derived from emotion versus neutral contrasts, meaning that, regardless of the task at hand, activation in these brain areas was greater when participants were experiencing or perceiving an emotion category than when they were experiencing or perceiving in a neutral control state. Activity in these brain regions is therefore integral to producing instances of emotion.

Our findings suggested the need to refine and add additional psychological operations to our model. Just as executive attention has been parsed into a set of distinguishable networks (e.g., Corbetta & Shulman Reference Corbetta and Shulman2002; Corbetta et al. Reference Corbetta, Patel and Shulman2008; Dosenbach et al. 2007; Seeley et al. Reference Seeley, Menon, Schatzberg, Keller, Glover, Kenna, Reiss and Greicius2007), we might further refine core affect into a set of smaller networks that correspond to even more basic mechanisms. For example, we might find separable networks corresponding to approach versus avoidance-related states. Our findings hint that brain regions in the left PFC might be candidates for a network involved in approach motivation because regions in the left lateral PFC (including the anterior and mid-insula, VLPFC, DLPFC and OFC) were consistently observed during instances of the experience of anger. This hypothesis is consistent with a large body of EEG evidence associating the left PFC with the experience of instances of anger (Harmon-Jones & Allen 1997; Harmon-Jones & Sigelman Reference Harmon-Jones and Sigelman2001) and approach motivation more generally (Amodio, et al. Reference Amodio, Master, Yee and Taylor2008; Fox Reference Fox1991; Sutton & Davidson Reference Sutton and Davidson1997). Future meta-analytic investigations should investigate the degree to which the left PFC and subcortical regions supporting incentive salience (e.g., ventral tegmentum, amygdala, and aspects of the nucleus accumbens and ventral pallidum; see Berridge & Robinson 2003) comprise a network for approach-related affect within the operation of core affect. We might also add ingredients for processing exteroceptive sensory sensations to our theoretical framework since visual cortex was one of the most frequently activated brain regions in our meta-analysis of discrete emotions. Exteroceptive sensory sensations are also important components of other types of mental states (e.g., perception, memory, judgments).

6.1. Alternate interpretations

Of course, there are alternate explanations for why we did not find strong evidence in support of a locationist framework. First, it is possible that neuroimaging is not well suited to yield evidence for functional specialization for emotion in the brain because of its spatial limitations. It therefore remains possible that scientists will find functional specialization for emotion at a more refined level of spatial analysis (e.g., at the level of smaller circuits or even cortical columns of neurons). While this always remains a possibility, it is important to note that even the most highly specialized brain regions (e.g., primary visual cortex) contain neurons that participate in different neural assemblies associated with different functions (e.g., Basole et al. Reference Basole, White and Fitzpatrick2003). This makes strong locationist interpretations of brain function unlikely when such interpretations attempt to find specificity for psychological categories (particularly at the level of the cortical column). Instead, the idea of neural re-use (e.g., Anderson Reference Anderson2010) is consistent with the psychological constructionist model of brain–emotion correspondence. Neuroimaging also has temporal limitations. It takes a few seconds for a BOLD response to materialize and we do not know whether emotions are episodes that extend over many seconds or are more instantaneous states that fire and resolve within that time frame. It is possible that scientists might find functional specialization for emotion at a more refined level of temporal analysis (e.g., using single cell recording), although such evidence has yet to be revealed. Moreover, we find it unlikely that neuroimaging and single cell recordings are measuring totally different phenomena. More likely, they are methods that complement one another (e.g., Horowitz 2005).

Second, it remains a possibility that we failed to locate a specific brain basis for discrete emotion categories because emotion categories are represented as anatomical networks of brain regions. Some researchers hypothesize that “resting state” analysesFootnote 10 of the brain's function, which reveal the intrinsic anatomical networks that chronically support the brain's fundamental processes, are influenced by anatomical connections (Deco et al. Reference Deco, Jirsa and McIntosh2011). If emotion categories were supported by anatomically given, inherited networks, then there should be intrinsic networks that correspond to the brain regions active during the experience or perception of instances of anger, sadness, fear, and other emotion categories. To date, however, no such intrinsic networks have been identified in broad inductive studies of such brain organization (e.g., Smith et al. Reference Smith, Fox, Miller, Glahn, Fox, Mackay, Filippini, Watkins, Toro, Laird and Beckman2009). Instead, the intrinsic networks that have thus far been identified bear resemblance to the psychological domains that are hypothesized by a psychological constructionist view (see functional groups in Kober et al. Reference Kober, Barrett, Joseph, Bliss-Moreau, Lindquist and Wager2008). For instance, the “default network” that is active when a person is not probed by an external stimulus in fMRI experiments (Buckner & Vincent Reference Buckner and Vincent2007; Raichle et al. Reference Raichle, MacLeod, Snyder, Powers, Gusnard and Shulman2001) is not only important to constructing representations of the past and the future, but also for constructing representations of emotion experience and perception in the moment. Portions of the intrinsic networks for “personal salience” (e.g., Seeley et al. Reference Seeley, Menon, Schatzberg, Keller, Glover, Kenna, Reiss and Greicius2007) and “executive control” (e.g., Corbetta & Shulman Reference Corbetta and Shulman2002; Seeley et al. Reference Seeley, Menon, Schatzberg, Keller, Glover, Kenna, Reiss and Greicius2007) appear anatomically similar to brain regions that we observed within our meta-analyses as well.

Of course, there is evidence for more limited brain circuits that correspond to specific behavioral adaptations in mammals. There is well-documented evidence for the anatomical circuitry underlying specific actions such as vocalizations (Jürgens Reference Jürgens2009), maternal behavior (e.g., pup retrieval, grooming, nest building, and nursing; Numan Reference Numan2007), freezing (Fanselow & Poulos Reference Fanselow and Poulos2005), startle (Davis et al. Reference Davis, Amaral and Winslow2008; Lang et al. Reference Lang, Davis and Öhman2000), attack (Blanchard & Blanchard Reference Blanchard and Blanchard2003) and appetitive behavior (Berridge & Kringelbach Reference Berridge and Kringelbach2008; Shultz Reference Shultz2006), to name just a few. In our view, these are just another set of basic operations and are not, in and of themselves, evidence that there is distinct anatomical circuitry for complex psychological categories such as sadness, love, fear, anger, or greed (each of which could contain instances of a range of behaviors) (Barrett et al. Reference Barrett, Lindquist, Bliss-Moreau, Duncan, Gendron, Mize and Brennan2007a). Animals produce actions in a way to maximize their outcome in a specific context, so many different behaviors can be associated with a given discrete emotion category; there is variety in the behaviors and the autonomics that populate any given emotion category. Sometimes, to make their findings more accessible, researchers will equate a specific behavior and its circuitry (e.g., freezing in the face of an uncertain danger) with an emotion category (e.g., fear). The problem with this logic is that it limits the definition of a complex psychological category to one or two behaviors. If fear is defined by freezing, then is fear not occurring at times when a rat flees, attacks, kicks bedding at a predator, or avoids an unknown corner of a maze? Or when humans avoid a dark alley, bungee jump, remember the events of September 11th, lock the door at night, or password-protect their bank accounts? As each of these actions is associated with a different neural network, which one is the fear network? If they are all fear networks, then what is the scientific value of the category fear for explaining behavior?

If discrete emotion categories are not associated with a specific brain locale, or even an anatomically inspired network that can be inherited, it is still possible that a pattern classification analysis on our meta-analytic database might reveal that each emotion category is represented by a specific combination of brain regions that co-activate together in time as a functional unit. We did not test this hypothesis, although we are in the process of developing these techniques for our software package. To the extent that such patterns are widely distributed across the brain, however (as opposed to being organized anatomically as inheritable units), such functionally defined networks for emotion categories (i.e., networks that only combine in a given context to produce a given type of mental state) would be consistent with a psychological constructionist (as opposed to a locationist) view.

It is possible that we failed to find evidence for the brain basis of discrete emotions because the methods employed in neuroimaging studies (or the laboratory for that matter) do not reliably elicit the type of discrete emotion experiences observed in the real world. After all, many scientists believe that emotions involve action (or action tendencies), and during scanning experiments participants must lie very still. Although this always remains a possibility, we do not believe it is a serious concern. First of all, even when participants are asked to lie still in a scanner, we still routinely observe increases in PAG activity in emotion. The PAG is necessary for motivated action patterns in animals, so these findings argue against the criticism that emotions invoked in the scanner are superficial. Even studies in which people are asked to imagine an emotional scenario probably create real experiences (as anyone knows who has become immersed in a mental reverie).

Finally, and perhaps most importantly, our observation that common brain activations exist across emotion categories is echoed in the pattern of findings for other (non-brain) measures of emotion. Since the beginning of psychology, researchers have questioned the idea that discrete emotion categories are each associated with a single, diagnostic pattern of response in the brain and body (e.g., Duffy Reference Duffy1934; Hunt Reference Hunt1941; James Reference James1884; for a review of such theories, see Gendron & Barrett Reference Gendron and Barrett2009). More recently, a number of empirical reviews (Barrett Reference Barrett2006a; Barrett et al. Reference Barrett, Lindquist, Bliss-Moreau, Duncan, Gendron, Mize and Brennan2007a; Mauss & Robinson Reference Mauss and Robinson2009; Ortony & Turner Reference Ortony and Turner1990; Russell Reference Russell2003) have highlighted the disconfirming evidence: Different discrete emotion categories are not distinguished by distinct patterns of peripheral physiology (Cacioppo et al. Reference Cacioppo, Berntson, Larsen, Poehlmann, Ito, Lewis and Haviland-Jones2000; Mauss & Robinson Reference Mauss and Robinson2009), facial muscle movements (Cacioppo et al. Reference Cacioppo, Berntson, Larsen, Poehlmann, Ito, Lewis and Haviland-Jones2000; Russell, Bachorowski & Fernandez-Dols Reference Russell, Bachorowski and Fernandez-Dols2003), vocal acoustics (Bachorowski & Owren Reference Bachorowski and Owren1995; Barrett Reference Barrett2006a; Russell, Bachorowski & Fernandez-Dols Reference Russell, Bachorowski and Fernandez-Dols2003) or by subcortical circuits in the mammalian brain (Barrett Reference Barrett2006a; Barrett et al. Reference Barrett, Lindquist, Bliss-Moreau, Duncan, Gendron, Mize and Brennan2007a). The present meta-analytic review adds to this literature by demonstrating that emotion categories do not map to discrete brain locales in the human brain either. Instead, evidence from our meta-analysis, as well as studies of emotion that use psychophysiological measures, objective measures of the face and voice, and subjective experience of emotion, all point to the idea that emotions emerge from a set of more basic operations (cf. Barrett Reference Barrett2006b). As neuroscientific methodologies progress, it will become increasingly important for scientists to formulate a viable conceptual framework for mapping emotions to the brain. Our findings suggest that a psychological constructionist approach might offer just such a framework. Locationist views might be deeply entrenched in commonsense, which makes the corresponding scientific models particularly compelling, but they do not match the scientific evidence in any measurement domain for emotion.

6.2. Future directions in the search for the brain basis of emotion

Despite the field's emphasis on locationist views (that inspired many of the experiments used in our meta-analysis), we found that the bulk of the empirical evidence is more consistent with the hypothesis that emotions emerge from the interplay of more basic psychological operations. We hypothesize that these operations and their corresponding neural networks influence and constrain one another to produce a variety of brain states that correspond to a variety of emotional states. To fully explore the power of a psychological constructionist approach in future research, researchers might combine traditional neuroimaging techniques with methods that make more network-based assumptions about brain function (e.g., Multivoxel Pattern Analysis: Haxby et al. Reference Haxby, Gobbini, Furey, Ishai, Schouten and Pietrini2001; Multivariate Partial Least Squares Analysis: McIntosh et al. Reference McIntosh, Bookstein, Haxby and Grady1996) (see Schienle & Schafer [Reference Schienle and Schäfer2009] for additional analysis approaches). Researchers might also utilize resting state analysis to identify functional networks that are intrinsic to the brain and compare those to the task-related assemblies of brain areas found across neuroimaging experiments (e.g., Smith et al. Reference Smith, Fox, Miller, Glahn, Fox, Mackay, Filippini, Watkins, Toro, Laird and Beckman2009). Finally, researchers must employ studies that capture and model the variability inherent in the collection of instances that form an emotion category like anger, disgust, fear, and so forth (e.g., Wilson-Mendenhall et al. Reference Wilson-Mendenhall, Barrett, Simmons and Barsalou2011). Most studies in our database utilize stimuli and induction techniques that invoke the most typical–and even caricatured–instances of an emotion category. Yet, daily experience tells us that there is great variability in the instances of anger, disgust, fear, happiness and sadness that we experience, and research bears this out. For example, an instance of disgust that occurs when watching others eat repulsive food involves a different brain state than an instance of disgust that occurs when watching surgical operations (Harrison et al. Reference Harrison, Gray, Giarnos and Critchley2010). Brain states that occur during instances of fear and anger are best described by an interaction between the content of the experience (e.g., whether the state is labeled fear vs. anger) and the context in which it occurred (e.g., a physical vs. social context) (Wilson-Mendenhall et al. Reference Wilson-Mendenhall, Barrett, Simmons and Barsalou2011).

7. Unifying the mind

A psychological constructionist approach is not only a viable approach for understanding the brain basis of emotion, but it might also offer a new psychological ontology for a neuroscientific approach to understanding the mind. If a psychological constructionist approach to the mind is correct, then some of psychology's time-honored folk distinctions become phenomenological distinctions. This has implications for understanding a range of psychological phenomena, including decision making, attention, visual perception, mental illness, and perhaps even consciousness more generally. Indeed, similar efforts are emerging in other psychological domains (Fuster Reference Fuster2006; Poldrack et al. Reference Poldrack, Halchenko and Hanson2009; Price & Friston Reference Price and Friston2005; Warnick et al. Reference Warnick, LaPorte and Kalueff2010). According to a psychological constructionist view of the mind, emotion does not influence cognition during decision-making as one pool ball exerts influence on another. Instead the view suggests that core affect, conceptualization, and executive attention (and perhaps other psychological operations) cooperate to realize a behavioral outcome. If this is the case, then we might not assume that emotion and cognition battle it out in the brain when a person makes the moral decision to sacrifice one life to save many (e.g., Greene et al. Reference Greene, Nystrom, Engell, Darley and Cohen2004), or that consumer decisions are predicated on competing affective and rational representations (e.g., Knutson et al. Reference Knutson, Rick, Wimmer, Prelec and Loewenstein2007). Instead, we might assume that affect and executive attention are merely different sources of attention in the brain, rather than processes that differ in kind (Barrett Reference Barrett2009b; Vuilleumier & Driver Reference Vuilleumier and Driver2007). Feeling and seeing might not be as distinct as is typically assumed (Barrett & Bar Reference Barrett and Bar2009; Duncan & Barrett Reference Duncan and Barrett2007). Even conceptions about “internal” versus “external” processing begin to break down when we take into account the fact that “internal” ingredients such as affect and conceptualization shape the very way in which exteroceptive sensory input is realized as perceptions by the brain (Bar 2009; Barrett & Bar Reference Barrett and Bar2009). A psychological constructionist framework of the mind thus begins to break down the most steadfast assumptions of our commonsense categories. In so doing, it charts a different but exciting path forward for the science of the mind.

SUPPLEMENTARY MATERIALS

Figure S1: http://www.journals.cambridge.org/bbs2012008

Figure S2: http://www.journals.cambridge.org/bbs2012008

Figure S3: http://www.journals.cambridge.org/bbs2012008

Figure S4: http://www.journals.cambridge.org/bbs2012008

Table S1: http://www.journals.cambridge.org/bbs2012008

Table S2: http://www.journals.cambridge.org/bbs2012008

Table S3: http://www.journals.cambridge.org/bbs2012008

Table S4: http://www.journals.cambridge.org/bbs2012008

Table S5: http://www.journals.cambridge.org/bbs2012008

Table S6: http://www.journals.cambridge.org/bbs2012008

Reference for supplementary material: http://www.journals.cambridge.org/bbs2012009

Prime elements of subjectively experienced feelings and desires: Imaging the emotional cocktail Buck, Ross W. Department of Communication Sciences and Psychology, University of Connecticut, Storrs, CT 06269-1085. http://coms.uconn.edu/directory/faculty/rbuck/index.htm Understanding emotion: Lessons from anxiety Button, Katherine S., Lewis, Glyn and Munafò, Marcus R. School of Social and Community Medicine, University of Bristol, Bristol, BS8 2BN, United Kingdom. http://www.bristol.ac.uk/psychiatry; School of Experimental Psychology, University of Bristol, Bristol BS8 1TU, United Kingdom. http://www.bristol.ac.uk/expsych/research/brain/targ Overcoming the emotion experience/expression dichotomy Caruana, Fausto and Gallese, Vittorio Department of Neuroscience, Section of Physiology, School of Medicine and Surgery, University of Parma, Università di Parma, 43100 Parma, Italy. http://www.unipr.it/arpa/mirror/english/staff/caruana.htm; Brain Center for Social and Motor Cognition, Italian Institute of Technology, 43100 Parma, Italy. http://www.unipr.it/arpa/mirror/english/staff/gallese.htm A constructionist account of emotional disorders Cramer, Angélique O. J., Kendler, Kenneth S. and Borsboom, Denny Department of Psychology, Faculty of Social and Behavioral Sciences, University of Amsterdam, 1018 WB Amsterdam, The Netherlands. / www.aojcramer.com http://sites.google.com/site/borsboomdenny/dennyborsboom; Department of Psychiatry, Virginia Commonwealth University, and Virginia Institute for Psychiatric and Genetics, Richmond, VA 23298-0126. Emotions as mind organs de Gelder, Beatrice and Vandenbulcke, Mathieu Cognitive and Affective Neuroscience Lab, Tilburg University, 5000 LE Tilburg, The Netherlands. www.beatricedegelder.com; Department of Neuroscience, Division of Psychiatry, Faculty of Medicine, University of Leuven, 3000 Leuven, Belgium. A rigorous approach for testing the constructionist hypotheses of brain function Deshpande, Gopikrishna, Sathian, K., Hu, Xiaoping and Buckhalt, Joseph A. Auburn University MRI Research Center, Department of Electrical and Computer Engineering, and Department of Psychology, Auburn University, Auburn, AL 36849. http://www.eng.auburn.edu/users/gzd0005/; Departments of Neurology, Rehabilitation Medicine, and Psychology, Emory University, and Atlanta VAMC Rehabilitation R&D Center of Excellence, Atlanta, GA 30322. http://neurology.emory.edu/Faculty/Sathian.htm; Coulter Department of Biomedical Engineering at Georgia Institute of Technology, and Center for Systems Imaging, Emory University, Atlanta, GA 30322. http://www.bme.emory.edu/~xhu/; Department of Special Education, Rehabilitation and Counseling, College of Education, Auburn University, Auburn, AL 36849-5222. http://www.auburn.edu/~buckhja/ Emotional participation in musical and non-musical behaviors Gardiner, Martin Frederick Center for the Study of Human Development, Brown University, Providence, RI 02912. What can neuroimaging meta-analyses really tell us about the nature of emotion? Hamann, Stephan Department of Psychology, Emory University, Atlanta, GA 30322. http://hamann.weebly.com/ Psychological constructionism and cultural neuroscience Hechtman, Lisa A., Pornpattananangkul, Narun and Chiao, Joan Y. Department of Psychology, Northwestern University, Evanston, IL 60208. http://culturalneuro.psych.northwestern.edu/Lab_Website/Welcome.html; Northwestern Interdepartmental Neuroscience Program, Evanston, IL 60208. Further routes to psychological constructionism Humeny, Courtney, Kelly, Deirdre and Brook, Andrew Institute of Cognitive Science, Carleton University, Ottawa, Ontario K1S 5B6, Canada. www.carleton.ca/~abrook Scaffolding emotions and evolving language Jablonka, Eva and Ginsburg, Simona Cohn Institute for the History and Philosophy of Science and Ideas, Tel Aviv University, Tel Aviv 69978, Israel. ; Department of Natural Science, The Open University of Israel, Raanana 43107, Israel. The sleeping brain and the neural basis of emotions Kirov, Roumen, Brand, Serge, Kolev, Vasil and Yordanova, Juliana Institute of Neurobiology, Bulgarian Academy of Sciences, 1113 Sofia, Bulgaria. http://www.bio.bas.bg/neurobiology/ http://www.bio.bas.bg/~cneurodyn/ http://www.bio.bas.bg/~cneurodyn/; Psychiatric Hospital of the University of Basel, 4012 Basel, Switzerland. http://www.upkbs.ch Emotion and personality factors influence the neural response to emotional stimuli Murphy, Fionnuala C., Ewbank, Michael P. and Calder, Andrew J. Medical Research Council, Cognition and Brain Sciences Unit, Cambridge CB2 7EF, United Kingdom. http://www.mrc-cbu.cam.ac.uk/ http://www.mrc-cbu.cam.ac.uk/ http://www.mrc-cbu.cam.ac.uk/ Emotions of “higher” cognitionFootnote 1 Perlovsky, Leonid Harvard University, Athinoula A. Martinos Center for Biomedical Imaging, Charlestown, MA 02129; and the Air Force Research Laboratory WPAFB, OH 45433. http://leonid-perlovsky.com/ Beyond brain regions: Network perspective of cognition–emotion interactions Pessoa, Luiz Department of Psychology, University of Maryland, College Park, MD 20742. http://emotioncognition.org The construction of emotional experience requires the integration of implicit and explicit emotional processes Quirin, Markus and Lane, Richard D. Institute of Psychology, University of Osnabrueck, 49074 Osnabrueck, Germany. http://www.motivationlab.uni-osnabrueck.de; Department of Psychiatry, University of Arizona, Tucson, AZ 85724-5002. http://www.psychiatry.arizona.edu/faculty/richard-d-lane-md-phd A systems approach to the brain basis of emotion also needs developmental and locationist views – the case of Tourette's Syndrome Rothenberger, Aribert Department of Child and Adolescent Psychiatry, University of Goettingen, D-37075 Göttingen, Germany. http://wwwuser.gwdg.de/~ukyk/ The role of the amygdala in the appraising brain Sander, David Laboratory for the Study of Emotion Elicitation and Expression, Department of Psychology, FPSE and Swiss Center for Affective Sciences, University of Geneva, CH-1205 Geneva, Switzerland. http://www.unige.ch/fapse/EmotionLab http://www.affective-sciences.org/ Functional specialization does not require a one-to-one mapping between brain regions and emotions Scarantino, Andrea Department of Philosophy and Neuroscience Institute, Georgia State University, Atlanta, GA 30302. https://sites.google.com/site/andreascarantinoswebsite/ Neuroscience findings are consistent with appraisal theories of emotion; but does the brain “respect” constructionism? Scherer, Klaus R. Swiss Center for Affective Sciences (SCAS), University of Geneva, CH-1205 Geneva, Switzerland. http://www.affective-sciences.org/user/scherer Invariants of human emotion Smaldino, Paul E. and Schank, Jeffrey C. Department of Psychology, University of California–Davis, Davis, CA 95616. http://psychology.ucdavis.edu/Grads/pesmaldino/ http://psychology.ucdavis.edu/faculty/Schank/ A rapprochement between emotion and cognition: Amygdala, emotion, and self-relevance in episodic-autobiographical memory Staniloiu, Angelica and Markowitsch, Hans J. Department of Physiological Psychology, University of Bielefeld, 33501 Bielefeld, Germany. http://uni-bielefeld.de/psychologie/ae/AE14/HOMEPAGE/Markowitsch_home.html; Centre for Addiction and Mental Health, Toronto, Ontario M5T 1R8, Canada. Feeling the strain: Predicting the third dimension of core affect Stapleton, Mog School of Philosophy, Psychology, and Language Sciences, University of Edinburgh, Edinburgh, Scotland EH8 9AD, United Kingdom. http://edinburgh.academia.edu/MogStapleton What's in a baby-cry? Locationist and constructionist frameworks in parental brain responses Swain, James E. and Ho, S. Shaun Department of Psychiatry, University of Michigan School of Medicine, Ann Arbor, MI 48109. http://myprofile.cos.com/jameseswain Narrative constructions and the life history issue in brain–emotions relations Unoka, Zsolt, Berán, Eszter and Pléh, Csaba Department of Psychiatry and Psychotherapy, Semmelweis University, 1083 Budapest, Hungary. ; Department of Psychology, Péter Pázmány Catholic University, 2087 Piliscsaba, Hungary. ; Department of Cognitive Science, Budapest University of Technology and Economics, 1111 Budapest, Hungary, and Department of Cognitive Science, Central European University, Budapest, Hungary. www.plehcsaba.hu Neuronal deactivation is equally important for understanding emotional processing Vigil, Jacob M., Dukes, Amber and Coulombe, Patrick Department of Psychology, University of New Mexico, Albuquerque, NM 87131-1161. http://www.unm.edu/~psych/faculty/sm_vigil.html Timing: A missing key ingredient in typical fMRI studies of emotion Waugh, Christian E. and Schirillo, James A. Department of Psychology, Wake Forest University, Winston-Salem, NC 27109-7778. emolab.psych.wfu.edu Need for more evolutionary and developmental perspective on basic emotional mechanisms Weisfeld, Glenn and LaFreniere, Peter Department of Psychology, Wayne State University, Detroit, MI 48202. www.clas.wayne.edu/unit-faculty-detail.asp?FacultyID=402; Department of Psychology, University of Maine, Orono, ME 04469. What are emotions and how are they created in the brain? Lindquist, Kristen A., Wager, Tor D., Bliss-Moreau, Eliza, Kober, Hedy and Barrett, Lisa Feldman Department of Neurology, Harvard Medical School/Massachusetts General Hospital/Martinos Center for Biomedical Imaging, Charlestown, MA 02129. http://www.nmr.mgh.harvard.edu/~lindqukr/; Department of Psychology, Harvard University, Cambridge, MA 02138; Department of Psychology and Neuroscience, University of Colorado, Boulder, CO 80309. http://www.psych.colorado.edu/~tor/; California National Primate Research Center, University of California, Davis, CA 95616. http://www.elizablissmoreau.com/; Department of Psychiatry and Behavioral Science, University of California, Davis, CA 95616; Department of Psychiatry, Yale University School of Medicine, New Haven, CT 06519. http://medicine.yale.edu/psychiatry/people/hedy_kober.profile; Department of Psychology, Northeastern University, Boston, MA 02115. http://www.affective-science.org/; Departments of Radiology and Psychiatry, Harvard Medical School/Massachusetts General Hospital/Martinos Center for Biomedical Imaging, Charlestown, MA 02129

ACKNOWLEDGMENTS

We thank Amitai Shenhav for his insights into the amygdala's the role in uncertainty, Natasha Sarkisian for her advice on logistic regression, and Kurt Gray for his help with Figure 5. We also thank James Russell, Elizabeth Kensinger, Hiram Brownell, Jerry Clore, and Stephan Hamann who commented on an earlier draft of this manuscript. This project was supported by a National Science Foundation (NSF) Graduate Research Fellowship and a Harvard University Mind/Brain/Behavior Initiative Postdoctoral Fellowship to Kristen Lindquist; a NSF Graduate Research Fellowship to Hedy Kober; a NSF grant (NSF 0631637) and a National Institute of Mental Health (NIMH) grant (R01MH076136) to Tor Wager; and a National Institutes of Health (NIH) Director's Pioneer Award (DP1OD003312), an National Institute on Aging (NIA) grant (R01 AG030311), and an Army Reasearch Institute (ARI) contract (W91WAW-08-C-0018) to Lisa Feldman Barrett. The views, opinions, and/or findings contained in this article are solely those of the authors and should not be construed as an official Department of the Army or Department of Defense position, policy, or decision.

Footnotes

1. These hypotheses have been inspired, in large part, by behavioral neuroscience research in nonhuman animals that has carefully mapped the circuitry for behavioral adaptations that occur in response to specific environmental challenges (e.g., freezing, attack, vocalizations). One variant of a locationist view focuses on the circuitry for behavioral adaptations such as freezing, escaping, aggressing, and so on (e.g., LeDoux Reference LeDoux2007; Panksepp Reference Panksepp1998), and assumes that one behavioral adaptation is at the core of each discrete emotion category. However, this one-to-one correspondence between a behavioral adaptation and a discrete emotion category has been challenged on the basis of existing research showing that mammals such as rats display a variety of behaviors based on what is most effective in a given context (for discussion, Barrett Reference Barrett2009a; Barrett et al. Reference Barrett, Lindquist, Bliss-Moreau, Duncan, Gendron, Mize and Brennan2007a).

2. For example, because the neurons within the amygdala are part of the neural reference space for discrete emotion, we can say with some certainly that the amygdala is likely to have increased activation when a person is experiencing or perceiving any emotion. This does not mean that the amygdala is necessary to each and every instance of emotion or even that it is specific to emotion, however. These ideas distinguish our approach from locationist accounts that assume that neurons within a given brain area (e.g., the amygdala) are consistently and specifically linked to a particular category of mental state (e.g., “fear”).

3. Here we use the term “functionally selective” to mean that a brain area can have some preference for certain mental states, even if it is not specific to that mental state. Functional selectivity might occur because a brain area supports a more basic psychological operation that helps to construct a certain mental state (e.g., the amygdala supports detection of salient exteroceptive sensations and is functionally selective for instances of fear). Functional selectivity does not refer to specificity, however. A brain area might be functionally selective for one mental state or even one basic psychological operation in one instance, and another state or operation in another instance. Functional selectivity is distinct from the concept of “selective influence” (cf. Sternberg Reference Sternberg2001), where a brain area being involved in one mental state (e.g., an instance of fear) but not another (e.g., an instance of anger) is taken as evidence of modularity.

4. For example, given that there is an increase in activation in the amygdala, the probability that a person is experiencing fear might be 0.7. The probability that he or she is experiencing another emotion (e.g., anger, disgust, happiness or sadness) is 1−0.7=0.3. The odds ratio=0.7/0.3=2.33. This means that given increased amygdala activation, the odds are 2.33 to 1 that the person is experiencing fear. In this case, the experience of fear is 113% more likely to predict increased activation in the amygdala than any other emotion state.

5. These findings might explain the amgydala's role in “fear learning” without assuming that the amygdala is specific to fear. In “fear learning,” for example, amygdala activity reflects orienting responses that occur when an organism learns to associate a neutral stimulus with an already salient stimulus. The amygdala contributes to the production of the skin conductance responses (SCRs) (Laine et al. Reference Laine, Spitler, Mosher and Gothard2009) used to index “fear learning.” Amygdala responses are associated with SCRs that occur immediately following the onset of a conditioned stimulus, suggesting that the amygdala is particularly involved in attention during learning but perhaps not the formation of associations (Cheng et al. Reference Cheng, Richards and Helmstetter2007; also see Blakeslee [Reference Blakeslee1979] and Spinks et al. [Reference Spinks, Blowers and Shek1985] for evidence that SCRs covary with changes in attention). This orienting account would also explain why increased amygdala activity is observed when animals learn to associate neutral stimuli with rewarding outcomes (e.g., Paton et al. Reference Paton, Belova, Morrison and Salzman2006; for a review see Murray Reference Murray2007), why amygdala activity corresponds to evaluative goals in the presence of both positive and negative stimuli (e.g., Cunningham et al. Reference Cunningham, Van Bavel and Johnsen2008; Paton et al. Reference Paton, Belova, Morrison and Salzman2006), and why stimulation of the amygdala facilitates orienting responses such as startle (Rosen & Davis Reference Rosen and Davis1988). Together, these findings make it clear why the amygdala is so ubiquitously involved in mammalian social behavior (i.e., male and female sexual behavior, maternal behavior, aggression; see Newman Reference Newman1999).

6. More than 90% (53/57) of study contrasts assessing fear perception in our database used startled faces that are unfamiliar to college students (who are typically participants in neuroimaging studies of healthy samples) (Whalen et al. Reference Whalen, Shin, McInerney, Fischer, Wright and Rauch2001) and are highly arousing (e.g., Russell & Bullock Reference Russell and Bullock1986). Approximately 35% (15/43) of study contrasts assessing the experience of disgust presented participants with images that were novel (i.e., infrequently experienced in the industrialized world) and highly arousing (i.e., containing contamination, mutilated body parts, maggots, etc.).

7. EEG findings do not associate instances of anger with OFC specifically, probably because EEG does not easily pick up activity in the orbital sector.

8. The medial OFC (mOFC) and the sACC, which are more generally part of VMPFC, were part of the neural reference space and are reported in separate sections. Aspects of VMPFC that do not include mOFC/sACC were part of the neural reference space, but were not significant at the thresholds we report in this article.

9. In some theoretical treatments of emotion, emotion regulation is thought to be a separate psychological event from emotion generation, with distinctive neural correlates; in a psychological constructionist approach, however, the processes are the same because there is no conceptual distinction between generation and regulation (Gross & Barrett Reference Gross and Barrett2011).

10. “Resting state” or “default” networks are evidenced as correlations between low-frequency signals in fMRI data that are recorded when there is no external stimulus or task. These networks are thought to be intrinsic in the human brain. For a review of intrinsic networks and their function, see Deco et al. (Reference Deco, Jirsa and McIntosh2011).

References

Addis, D. R., Wong, A. T. & Schacter, D. L. (2007) Remembering the past and imagining the future: Common and distinct neural substrates during event construction and elaboration. Neuropsychologia 45:1363–77. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed.CrossRefGoogle ScholarPubMed
Adolphs, R. (2008) Fear, faces, and the human amygdala. Current Opinion in Neurobiology 18:166–72. Available at:http://www.ncbi.nlm.nih.gov/pubmed/18655833.CrossRefGoogle ScholarPubMed
Adolphs, R. (2009) The social brain: Neural basis of social knowledge. Annual Review of Psychology 60:693716. Available at:http://www.ncbi.nlm.nih.gov/pubmed/18771388.CrossRefGoogle ScholarPubMed
Adolphs, R. (2010) What does the amygdala contribute to social cognition? Annals of the New York Academy of Sciences: The year in Cognitive Neuroscience 1191(1):4261. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2871162/.CrossRefGoogle ScholarPubMed
Adolphs, R., Gosselin, F., Buchanan, T. W., Tranel, D., Schyns, P. & Damasio, A. R. (2005) A mechanism for impaired fear recognition after amygdala damage. Nature 433:6872. Available at:http://www.nature.com/nature/journal/v433/n7021/abs/nature03086.html.CrossRefGoogle ScholarPubMed
Adolphs, R. & Tranel, D. (1999) Intact recognition of emotional prosody following amygdala damage. Neuropsychologia 37:1285–92. Available at:http://emotion.caltech.edu/papers/AdolphsTranel1999Intact.pdf.CrossRefGoogle ScholarPubMed
Adolphs, R., Tranel, D. & Damasio, A. R. (2003) Dissociable neural systems for recognizing emotions. Brain and Cognition 52:6169.CrossRefGoogle ScholarPubMed
Adolphs, R., Tranel, D., Damasio, H. & Damasio, A. (1994) Impaired recognition of emotion in facial expressions following bilateral damage to the human amygdala. Nature 372:669–72. Available at:http://www.nature.com/doifinder/10.1038/372669a0.CrossRefGoogle Scholar
Adolphs, R., Tranel, D., Damasio, H. & Damasio, A. R. (1995) Fear and the human amygdala. Journal of Neuroscience 15:5879. Available at:http://www.jneurosci.org/cgi/content/abstract/15/9/5879.CrossRefGoogle ScholarPubMed
Adolphs, R., Tranel, D., Hamann, S., Young, A. W., Calder, A. J., Phelps, E. A., Anderson, A., Lee, G. P. & Damasio, A. R (1999) Recognition of facial emotion in nine individuals with bilateral amygdala damage. Neuropsychologia 37:1111–17. Available at:http://www.cs.phs.uoa.gr/el/courses/emotions/papers/ADOPLHS%201999.pdf.CrossRefGoogle ScholarPubMed
Amaral, D. G. & Price, J. L. (1984) Amygdalo-cortical projections in the monkey (Macaca fascicularis). Journal of Comparative Neurology 230:465–96. Available at:http://www3.interscience.wiley.com/journal/109688079/abstract.CrossRefGoogle ScholarPubMed
Amodio, D. M., Master, S. L., Yee, C. M. & Taylor, S. H. (2008) Neurocognitive components of the behavioral inhibition and activation systems. Implications for theories of self-regulation. Psychophysiology 45:1119.CrossRefGoogle ScholarPubMed
An, X., Bandler, R., Ongur, D. & Price, J. L. (1998) Prefrontal cortical projections to longitudinal columns in the midbrain periaqueductal gray in macaque monkeys. The Journal of Comparative Neurology 401:455–79. Available at; http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/9826273.3.0.CO;2-6>CrossRefGoogle ScholarPubMed
Anderson, E., Siegel, E. H., Bliss-Moreau, E. & Barrett, L. F. (2011). The visual impact of gossip. Science 332:1446–48. Available at:http://www.sciencemag.org/cgi/rapidpdf/science.1201574?ijkey=Y/jsFjEWqreKA&keytype=ref&siteid=sci CrossRefGoogle ScholarPubMed
Anderson, A. K. & Phelps, E. A. (1998) Intact recognition of vocal expressions of fear following bilateral lesions of the amygdala. NeuroReport 9:3607–13. Available at; http://www.ncbi.nlm.nih.gov/pubmed/9858368.Google Scholar
Anderson, A. K. & Phelps, E. A. (2001) Lesions of the human amygdala impair enhanced perception of emotionally salient events. Nature 411:305309. http://www.psych.nyu.edu/phelpslab/abstracts/lesions_amygdala.pdf.CrossRefGoogle ScholarPubMed
Anderson, A. K. & Phelps, E. A. (2002) Is the human amygdala critical for the subjective experience of emotion? Evidence of intact dispositional affect in patients with amygdala lesions. Journal of Cognitive Neuroscience 14:709–20. Available at:http://www.psych.nyu.edu/phelpslab/abstracts/anderson_phelps2002.pdf.CrossRefGoogle ScholarPubMed
Anderson, M. L. (2010) Neural reuse: A fundamental organizational principle of the brain. Behavioral and Brain Sciences 33:245313. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/20964882.CrossRefGoogle ScholarPubMed
Angyal, A. (1941) Disgust and related aversions. Journal of Abnormal and Social Psychology 36:393412.CrossRefGoogle Scholar
Aron, A. R., Robbins, T. W. & Poldrack, R. A. (2004) Inhibition and the right inferior frontal cortex. Trends in Cognitive Sciences 8:170–77. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/15050513.CrossRefGoogle ScholarPubMed
Atkinson, A. P., Heberlein, A. S. & Adolphs, R. (2007) Spared ability to recognise fear from static and moving whole-body cues following bilateral amygdala damage. Neuropsychologia 45:2772–82. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1988781/.CrossRefGoogle ScholarPubMed
Bach, D. R., Schachinger, H., Neuhoff, J. G., Esposito, F. Di Salle, F., Lehmann, C., Herdener, M., Scheffler, K. & Seifrit, E. (2008) Rising sound intensity: An intrinsic warning cue activating the amygdala. Cerebral Cortex 18: 15150. Available at:http://cercor.oxfordjournals.org/content/18/1/145.full CrossRefGoogle ScholarPubMed
Bachorowski, J. A. & Owren, M. J. (1995) Vocal expression of emotion: Acoustic properties of speech are associated with emotional intensity and context. Psychological Science 6:219–24. Available at:http://pss.sagepub.com/content/6/4/219.abstract.CrossRefGoogle Scholar
Badre, D. & Wagner, A. D. (2007) Left ventrolateral prefrontal cortex and the cognitive control of memory. Neuropsychologia 45:2883–901. Available at:http://www.stanford.edu/group/memorylab/.CrossRefGoogle ScholarPubMed
Bancaud, J., Brunet-Bourgin, F., Chauvel, P. & Halgren, E. (1994) Anatomical origin of deja vu and vivid “memories” in human temporal lobe epilepsy. Brain 117:7190. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/8149215.CrossRefGoogle Scholar
Banich, M. T. (2009) Executive function: The search for an integrated account. Current Directions in Psychological Science 18(2):8994. Available at:http://psych.colorado.edu/~mbanich.CrossRefGoogle Scholar
Bar, M. (2007) The proactive brain: Using analogies and associations to generate predictions. Trends in Cognitive Sciences 11:280–89.CrossRefGoogle ScholarPubMed
Bar, M. (2009b) The proactive brain: Memory for predictions. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 364:1235–43. Available at:http://rstb.royalsocietypublishing.org/content/364/1521/1235.abstract.CrossRefGoogle ScholarPubMed
Bar, M., Kassam, K. S., Ghuman, A. S., Boshyan, J., Schmid, A. M., Dale, A. M., Hämäläinen, M. S., Marinkovic, K., Schacter, D. L. & Rosen, B. R. (2006) Top-down facilitation of visual recognition. Proceedings of the National Academy of Sciences 103:449–54. Available at:http://www.pnas.org/content/103/2/449.full.CrossRefGoogle ScholarPubMed
Barbas, H. (1988) Anatomic organization of basoventral and mediodorsal visual recipient prefrontal regions in the rhesus monkey. The Journal of Comparative Neurology 276:313–42. Available at:http://www3.interscience.wiley.com/journal/109690199/abstract.CrossRefGoogle ScholarPubMed
Barbas, H., Henion, T. H. & Dermon, C. R. (1991) Diverse thalamic projections to the prefrontal cortex in the rhesus monkey. The Journal of Comparative Neurology 313:6594. Available at:http://www3.interscience.wiley.com/journal/109692035/abstract.CrossRefGoogle Scholar
Barbas, H. & Pandya, D. N. (1989) Architecture and intrinsic connections of the prefrontal cortex in the rhesus monkey. The Journal of Comparative Neurology 286:353–75. Available at:http://www3.interscience.wiley.com/journal/109690672/abstract.CrossRefGoogle ScholarPubMed
Barrett, L. F. (2006a) Are emotions natural kinds? Perspectives on Psychological Science 1(1):2858. Available at:http://www.bc.edu/sites/asi/publications/lfb/Barrett2006kinds.pdf.CrossRefGoogle ScholarPubMed
Barrett, L. F. (2006b) Solving the emotion paradox: Categorization and the experience of emotion. Personality and Social Psychology Review 10:2046. Available at:http://psr.sagepub.com/cgi/content/abstract/10/1/20.CrossRefGoogle ScholarPubMed
Barrett, L. F. (2009a) The future of psychology: Connecting mind to brain. Perspectives on Psychological Science 4:326–39. Available at:http://www.affective-science.org/pubs/2009/barrett2009-future-psych.pdf.CrossRefGoogle Scholar
Barrett, L. F. (2009b) Variety is the spice of life: A psychological construction approach to understanding variability in emotion. Cognition and Emotion 23:1284–306. Available at:http://www.affective-science.org/pubs/2009/variety-2009.pdf.CrossRefGoogle ScholarPubMed
Barrett, L. F. (2011a) Bridging token identity theory and supervenience theory through psychological construction. Psychological Inquiry 22:115–27.CrossRefGoogle ScholarPubMed
Barrett, L. F. & Bar, M. (2009) See it with feeling: Affective predictions in the human brain. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences 364:1325–34. Available at:http://barlab.mgh.harvard.edu/papers/RSBarrettBar.pdf.CrossRefGoogle Scholar
Barrett, L. F. & Bliss-Moreau, E. (2009) Affect as a psychological primitive. Advances in Experimental Social Psychology 41:167218. Available at:http://www3.interscience.wiley.com/journal/122501435/abstract.CrossRefGoogle ScholarPubMed
Barrett, L. F. & Kensinger, E. A. (2010) Context is routinely encoded during emotion perception. Psychological Science 21(4):595–99.CrossRefGoogle ScholarPubMed
Barrett, L. F., Lindquist, K. A., Bliss-Moreau, E., Duncan, S., Gendron, M., Mize, J. & Brennan, L. (2007a) Of mice and men: Natural kinds of emotions in the mammalian brain? A response to Panksepp and Izard. Perspectives on Psychological Science 2(3):297311. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2597798/.CrossRefGoogle Scholar
Barrett, L. F., Lindquist, K. A. & Gendron, M. (2007b) Language as context for the perception of emotion. Trends in Cognitive Sciences 11:327–32. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2225544/.CrossRefGoogle ScholarPubMed
Barrett, L. F., Mesquita, B., Ochsner, K. N. & Gross, J. J. (2007c) The experience of emotion. Annual Review of Psychology 58:373403. Available at:http://arjournals.annualreviews.org/doi/full/10.1146/annurev.psych.58.110405.085709.CrossRefGoogle ScholarPubMed
Barrett, L. F., Ochsner, K. N. & Gross, J. J. (2007d) On the automaticity of emotion. In: Social psychology and the unconscious: The automaticity of higher mental processes, ed. Bargh, J., pp. 173217. Psychology Press.Google Scholar
Barrett, L. F., Tugade, M. M. & Engle, R. W. (2004) Individual differences in working memory capacity and dual-process theories of the mind. Psychological Bulletin 130:553–73. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1351135/ CrossRefGoogle ScholarPubMed
Barrett, L. F. & Wager, T. D. (2006) The structure of emotion: Evidence from neuroimaging studies. Current Directions in Psychological Science 15:79–83. Available at:http://psycnet.apa.org/index.cfm?fa=search.displayRecord&uid=2006-06699-007.CrossRefGoogle Scholar
Barsalou, L. W. (2003) Situated simulation in the human conceptual system. Language and Cognitive Processes 18:513–62.CrossRefGoogle Scholar
Barsalou, L. W. & Wiemer-Hastings, K. (2005) Situating abstract concepts. In: Grounding cognition: The role of perception and action in memory, language, and thought, ed. Pecher, D. & Zwaan, R., pp. 129–63. Cambridge University Press. Available at:http://psychology.emory.edu/cognition/barsalou/papers/Barsalou_Wiemer-Hastings_chap_2005_abstract_concepts.pdf.CrossRefGoogle Scholar
Basole, A., White, L. E. & Fitzpatrick, D. (2003) Mapping multiple features in the population response of visual cortex. Nature 423:986–90. Available at:http://www.nature.com/nature/journal/v423/n6943/abs/nature01721.html.CrossRefGoogle ScholarPubMed
Bechara, A., Tranel, D. & Damasio, H. (2000) Characterization of the decision-making deficit of patients with ventromedial prefrontal cortex lesions. Brain: A Journal of Neurology 123:2189–202. Available at:http://brain.oxfordjournals.org/cgi/content/abstract/123/11/2189.CrossRefGoogle ScholarPubMed
Bechara, A., Tranel, D., Damasio, H., Adolphs, R., Rockland, C. & Damasio, A. (1995) Double dissociation of conditioning and declarative knowledge relative to the amygdala and hippocampus in humans. Science 269:1115–18. Available at:http://www.sciencemag.org/cgi/content/abstract/sci;269/5227/1115.CrossRefGoogle Scholar
Bechara, A., Tranel, D., Damasio, H. & Damasio, A. R. (1996) Failure to respond autonomically to anticipated future outcomes following damage to prefrontal cortex. Cerebral Cortex 6:215–25. Available at:http://cercor.oxfordjournals.org/cgi/content/abstract/6/2/215.CrossRefGoogle ScholarPubMed
Bechtel, W. (2008) Mental mechanisms. Philosophical perspectives on cognitive neuroscience. Taylor and Francis. Available at:http://www.amazon.com/Mental-Mechanisms-Philosophical-Perspectives-Neuroscience/dp/0805863346/ref=sr_1_3?s=books&ie=UTF8&qid=1296488925&sr=1-3.Google Scholar
Beer, J. S., John, O., Scabini, D., Knight, R. T. (2003) Orbitofrontal cortex and social behavior: Integrating self-monitoring and emotion–cognition interactions. Journal of Cognitive Neuroscience 18:871–79. Available at:http://www.ncbi.nlm.nih.gov/pubmed/14561114.CrossRefGoogle Scholar
Behbehani, M. M. (1995) Functional characteristics of the midbrain periaqueductal gray. Progress in Neurobiology 46:575605. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/8545545.CrossRefGoogle ScholarPubMed
Berridge, K. C. & Kringelbach, M. L. (2008) Affective neuroscience of pleasure: Reward in humans and animals. Psychopharmacology 199:457–80. Available at:http://www.springerlink.com/content/h10384324x566083/.CrossRefGoogle ScholarPubMed
Blackford, J. U., Buckholtz, J. W., Avery, S. N. & Zald, D. H. (2010) A unique role for human amygdala in novelty detection. Neuroimage 50:1188–93. Available at:http://www.ncbi.nlm.nih.gov/pubmed/20045069.CrossRefGoogle ScholarPubMed
Blakeslee, P. (1979) Attention and vigilance: Performance and skin conductance response changes. Psychophysiology 16:413–19. Available at:http://www3.interscience.wiley.com/journal/119602640/abstract.CrossRefGoogle ScholarPubMed
Blanchard, D. C. & Blanchard, R. J. (2003) What can animal aggression tell us about human aggression? Hormones and Behavior 44:171–77. Available at:http://www.ncbi.nlm.nih.gov/pubmed/14609539.CrossRefGoogle ScholarPubMed
Bliss-Moreau, E., Toscano, J. E., Baumann, M. D., Mason, W. A. & Amaral, D. G. (2010) Neonatal amygdala or hippocampus lesions influence responsiveness to objects. Developmental Psychobiology 52:487503. Available at:http://onlinelibrary.wiley.com/doi/10.1002/dev.20451/abstract.CrossRefGoogle ScholarPubMed
Blumberg, M. S. & Sokoloff, G. (2001) Do infant rats cry? Psychological Review 108:8395. Available at:http://psycnet.apa.org/journals/rev/108/1/83.html.CrossRefGoogle ScholarPubMed
Bradley, M. M., Codispoti, M., Cuthbert, B. N. & Lang, P. J. (2001) Emotion and motivation I: Defensive and appetitive reactions in picture processing. Emotion 1(3):276–98.CrossRefGoogle ScholarPubMed
Breiter, H. C., Etcoff, N. L., Whalen, P. J., Kennedy, W. A., Rauch, S. L., Buckner, R. L., Strauss, M. M., Hyman, S. E. & Rosen, B. R. (1996) Response and habituation of the human amygdala. Neuron 17:875–77. Available at:http://www.ncbi.nlm.nih.gov/pubmed/8938120.CrossRefGoogle ScholarPubMed
Brierley, B., Medford, N., Shaw, P. & David, A. S. (2004) Emotional memory and perception in temporal lobectomy patients with amygdala damage. Journal of Neurology, Neurosurgery, and Psychiatry 75:593–99. Available at:http://jnnp.bmj.com/content/75/4/593.abstract.CrossRefGoogle ScholarPubMed
Büchel, C., Dolan, R. J., Armony, J. L. & Friston, K. J. (1999) Amygdala-hippocampal involvement in human aversive trace conditioning revealed through event-related functional magnetic resonance imaging. Journal of Neuroscience 19:10869–76. Available at:http://www.ncbi.nlm.nih.gov/pubmed/10594068.CrossRefGoogle ScholarPubMed
Buckner, R. L., Andrews-Hanna, J. R. & Schacter, D. L. (2008) The brain's default network: Anatomy, function, and relevance to disease. Annals of the New York Academy of Sciences 1124:138. Available at:http://www.wjh.harvard.edu/~dsweb/pdfs/08_03_RLB_JRA_DLS.pdf.CrossRefGoogle ScholarPubMed
Buckner, R. L. & Carroll, D. C. (2007) Self-projection and the brain. Trends in Cognitive Sciences 11:4957. Available at:http://www.ncbi.nlm.nih.gov/pubmed/17188554.CrossRefGoogle ScholarPubMed
Buckner, R. L. & Vincent, J. L. (2007) Unrest at rest: Default activity and spontaneous network correlations. NeuroImage 37:1091–96. Available at:http://www.nmr.mgh.harvard.edu/nexus/publications/publications.html.CrossRefGoogle ScholarPubMed
Burns, L. H., Annett, L., Kelley, A. E., Everitt, B. J. & Robbins, T. W. (1996) Effects of lesions to amygdala, ventral subiculum, medial prefrontal cortex, and nucleus accumbens on the reaction to novelty: Implication for limbic-striatal interactions. Behavioral Neuroscience 110:6073. Available at:http://psycnet.apa.org/journals/bne/110/1/60.html.CrossRefGoogle ScholarPubMed
Butter, C. & Snyder, D. (1972) Alterations in aversive and aggressive behaviors following orbital frontal lesions in rhesus monkeys. Acta Neurobiolagiae Experimentalis 32:525–65. Available at:http://psycnet.apa.org/index.cfm?fa=search.displayRecord&uid=1973-08597-001.Google ScholarPubMed
Cabanac, M. (2002) What is emotion? Behavioural Processes 60:6983.CrossRefGoogle ScholarPubMed
Cacioppo, J. T., Berntson, C. G., Larsen, J. T., Poehlmann, K. M. & Ito, T. A. (2000) The psychophysiology of emotion. In: Handbook of emotions, 2nd edition, ed. Lewis, M. & Haviland-Jones, J. M., pp. 173–91. Guilford Press.Google Scholar
Cacioppo, J., Berntson, C. & Nusbaum, H. C. (2008) Neuroimaging as a new tool in the toolbox of psychological science. Current Directions in Psychological Science 17:6267. Available at:http://psychology.uchicago.edu/people/faculty/cacioppo/jtcreprints/cbn08.pdf.CrossRefGoogle Scholar
Calder, A. J. (2003) Disgust discussed. Annals of Neurology 53:427–28. Available at:http://www.ncbi.nlm.nih.gov/pubmed/12666109.CrossRefGoogle ScholarPubMed
Calder, A. J., Keane, J., Manes, F., Antoun, N. & Young, A. W. (2000) Impaired recognition and experience of disgust following brain injury. Nature Neuroscience 3:1077–78. Available at:http://psycnet.apa.org/index.cfm?fa=search.displayRecord&uid=2000-16375-003.CrossRefGoogle ScholarPubMed
Calder, A. J., Keane, J., Young, A. W., Lawrence, A. D., Mason, S. & Barker, R. A. (2010) The relation between anger and different forms of disgust. Implications for emotion recognition impairments in Huntington's disease. Neuropsychologia 48:2719–29.CrossRefGoogle ScholarPubMed
Calder, A. J., Lawrence, A. D. & Young, A. W. (2001) Neuropsychology of fear and loathing. Nature Reviews Neuroscience 2:352–63. Available at:http://jppsg.ac.uk/psych/resources/lawrence2001.pdf.CrossRefGoogle Scholar
Carmichael, S. T. & Price, J. L. (1995) Sensory and premotor connections of the orbital and medial prefrontal cortex of macaque monkeys. Journal of Comparative Neurology 363:642–64. Available at:http://www.ncbi.nlm.nih.gov/pubmed/8847422.CrossRefGoogle ScholarPubMed
Carrive, P., Bandler, R. & Dampney, R. A. (1989) Vicerotopic control of regional vascular beds by discrete groups of neurons within the midrain periaqueductal gray. Brain Research 493:385–90. Available at:http://www.ncbi.nlm.nih.gov/pubmed/2765906.CrossRefGoogle Scholar
Champod, A. S. & Petrides, M. (2007) Dissociable roles of the posterior parietal and the prefrontal cortex in manipulation and monitoring processes. Proceedings of the National Academy of Sciences USA 104:14837–42. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/17804811.CrossRefGoogle ScholarPubMed
Cheng, D. T., Richards, J. & Helmstetter, F. J. (2007) Activity in the human amygdala corresponds to early, rather than late period autonomic responses to a signal for shock. Learning and Memory 14:485–90. Available at:http://learnmem.cshlp.org/content/14/7/485.abstract.CrossRefGoogle ScholarPubMed
Chudasama, Y. & Robbins, T. W. (2003) Dissociable contributions of the orbitofrontal and infralimbic cortex to pavlovian autoshaping and discrimination reversal learning: Further evidence for the functional heterogeneity of the rodent frontal cortex. The Journal of Neuroscience 23:8771–80. Available at; http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/14507977.CrossRefGoogle ScholarPubMed
Clore, G. L. & Ortony, A. (2008) Appraisal theories: How cognition shapes affect into emotion. In: Handbook of emotions, 3rd edition, ed. Lewis, M., Haviland-Jones, J. M. & Barrett, L. F., pp. 628–42. Guilford Press.Google Scholar
Cole, M. W. & Schneider, W. (2007) The cognitive control network: Integrated cortical regions with dissociable functions. Neuroimage 37:343–60. Available at:http://www.ncbi.nlm.nih.gov/pubmed/17553704.CrossRefGoogle ScholarPubMed
Constantinidis, C., Williams, C. V. & Goldman-Rakic, P. S. (2002) A role for inhibition in shaping the temporal flow of information in prefrontal cortex. Nature Neuroscience 5:175–80. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/11802172.CrossRefGoogle ScholarPubMed
Corbetta, M., Patel, G. & Shulman, G. L. (2008) The reorienting system of the human brain: From environment to theory of mind. Neuron 58:306–24. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2441869/.CrossRefGoogle ScholarPubMed
Corbetta, M. & Shulman, G. L. (2002) Control of goal-directed and stimulus-driven attention in the brain. Nature Reviews Neuroscience 3:215–29. Available at:http://www.nature.com/nrn/journal/v3/n3/execsumm/nrn755.html.CrossRefGoogle ScholarPubMed
Costafreda, S. G., Brammer, M. J., David, A. S. & Fu, C. H. Y. (2008) Predictors of amygdala activation during the processing of emotional stimuli: A meta-analysis of 385 PET and fMRI studies. Brain Research Reviews 58:5770. Available at:http://www.sciencedirect.com/science/article/pii/S0165017307002482 CrossRefGoogle ScholarPubMed
Coyne, J. C. (1994) Self-reported distress: Analog or ersatz depression? Psychological Bulletin 116(1):2945. Available at:http://psycnet.apa.org/journals/bul/116/1/29.html.CrossRefGoogle ScholarPubMed
Craig, A. D. (2002) How do you feel? Interoception: The sense of the physiological condition of the body. Nature Reviews Neuroscience 3:655–66. Available at:http://www.nature.com/nrn/journal/v3/n8/box/nrn894_BX1.html.CrossRefGoogle Scholar
Craig, A. D. B. (2009) How do you feel–now? The anterior insula and human awareness. Nature Reviews. Neuroscience 10:5970. Available at:http://www.nature.com/nrn/journal/v10/n1/full/nrn2555.html.CrossRefGoogle Scholar
Critchley, H. D., Mathias, C. J. & Dolan, R. J. (2001) Neuroanatomical basis for first and second order representations of bodily states. Nature Neuroscience 4:207–12. Available at:http://www.ncbi.nlm.nih.gov/pubmed/11175883.CrossRefGoogle ScholarPubMed
Cunningham, W. A., Van Bavel, J. J. & Johnsen, I. R. (2008) Affective flexibility. Evaluative processing goals shape amygdala activity. Psychological Science 19:152–60. Available at:http://faculty.psy.ohio-state.edu/cunningham/pdf/2008_Cunningham_etal_PS.pdf.CrossRefGoogle ScholarPubMed
Curtis, V., Aunger, R. & Rabie, T. (2004) Evidence that disgust evolved to protect from risk of disease. Proceedings of the Royal Society of London, Series B: Biological Sciences 271:131–33. Available at:http://rspb.royalsocietypublishing.org/content/271/Suppl_4/S131.full.pdf.CrossRefGoogle ScholarPubMed
Damasio, A. R. (1999) The feeling of what happens: Body and emotion in the making of consciousness. Harcourt Brace/Random House.Google Scholar
Damaraju, E., Huang, Y-M., Barrett, L. F. & Pessoa, L. (2009). Affective learning enhances activity and functional connectivity in early visual cortex. Neuropsychologia 47:2480–87.CrossRefGoogle Scholar
Damasio, A., Tranel, D. & Damasio, H. (1990) Individuals with sociopathic behavior caused by frontal damage fail to respond autonomically to social stimuli. Behavioral Brain Research 41:8194. Available at:http://www.ncbi.nlm.nih.gov/pubmed/2288668.CrossRefGoogle ScholarPubMed
Damsa, C., Kosel, M. & Moussally, J. (2009) Current status of brain imaging in anxiety disorders. Current Opinion in Psychiatry 22:96110. Available at:http://journals.lww.com/co-psychiatry/Abstract/2009/01000/Current_status_of_brain_imaging_in_anxiety.17.aspx.CrossRefGoogle ScholarPubMed
Davis, M. (1992) The role of the amygdala in fear and anxiety. Annual Review of Neuroscience 15:353–75. Available at:http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.ne.15.030192.002033.CrossRefGoogle ScholarPubMed
Davis, M. Antoniadis, E., Amaral, D. G. & Winslow, J. T. (2008) Acoustic startle reflex in monkeys: A review. Reviews in the Neurosciences 19:171–85. Available at:http://www.ncbi.nlm.nih.gov/pubmed/18751523.CrossRefGoogle ScholarPubMed
de Bruin, J. P. (1990) Social behavior in the prefrontal cortex. Progress in Brain Research 85:495–96. Available at:http://www.ncbi.nlm.nih.gov/pubmed/2094911.Google ScholarPubMed
de Bruin, J. P., Van Oyen, H. G. & Van De Poll, N. (1983) Behavioural changes following lesions of the orbital prefrontal cortex in male rats. Behavioural Brain Research 10:209–32. Available at:http://www.ncbi.nlm.nih.gov/pubmed/6686460.CrossRefGoogle ScholarPubMed
Deco, G., Jirsa, V. K. & McIntosh, A. R. (2011) Emerging concepts for the dynamical organization of resting-state activity in the brain. Nature Reviews Neuroscience 12:4356. Available at:http://www.ncbi.nlm.nih.gov/pubmed/21170073.CrossRefGoogle ScholarPubMed
Deets, A. C., Harlow, H. F., Singh, S. D. & Blomquist, A. J. (1970) Effects of bilateral lesions of the frontal granular cortex on the social behavior of rhesus monkeys. Journal of Comparative Physiological Psychology 72:452–61. Available at:http://www.ncbi.nlm.nih.gov/sites/entrez.CrossRefGoogle ScholarPubMed
Devinsky, O., Morrell, M. J. & Vogt, B. A. (1995) Contributions of anterior cingulate cortex to behaviour. Brain 118:279. Available at:http://brain.oxfordjournals.org/cgi/content/abstract/118/1/279.CrossRefGoogle ScholarPubMed
Dewar, K. & Xu, F. (2009) Do early nouns refer to kinds or distinct shapes? Evidence from 10-month-old infants. Psychological Science 20:252–57.CrossRefGoogle ScholarPubMed
Dosenbach, N. U., Visscher, K. M., Palmer, E. D., Miezin, F. M., Wenger, K. K., Kang, H. C., Burgund, E. D., Grimes, A. L., Schlaggar, B. L. & Petersen, S. E. (2006) A core system for the implementation of task sets. Neuron 50:799812. Available at:http://www.ncbi.nlm.nih.gov/pubmed/16731517.CrossRefGoogle ScholarPubMed
Duffy, E. (1934) Emotion: An example of the need for reorientation in psychology. Psychological Review 41:184–98. Available at:http://psycnet.apa.org/journals/rev/41/2/184/.CrossRefGoogle Scholar
Duffy, E. (1957) The psychological significance of the concept of “arousal” or “activation.” Psychological Review 64:265–75. Available at:http://psycnet.apa.org/journals/rev/64/5/265/.CrossRefGoogle ScholarPubMed
Duncan, S. & Barrett, L. F. (2007) Affect is a form of cognition: A neurobiological analysis. Cognition and Emotion 21(6):1184–211. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2396787/.CrossRefGoogle ScholarPubMed
Eblen, F. & Graybiel, A. M. (1995) Highly restricted origin of prefrontal cortical inputs to striosomes in the macaque monkey. Journal of Neuroscience 15:59996013. Available at:http://www.ncbi.nlm.nih.gov/sites/entrez.CrossRefGoogle ScholarPubMed
Eckert, M. A., Menon, V., Walczak, A., Ahlstrom, J., Denslow, S., Horowitz, A. & Dubno, J. R. (2009) At the heart of the ventral attention system: The right anterior insula. Human Brain Mapping 30:2530–41. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pmc/articles/PMC2712290/.CrossRefGoogle ScholarPubMed
Edelman, G. M. (1989) The remembered present: A biological theory of consciousness. MIT Press. Available at:http://www.amazon.com/Remembered-Present-Biological-Theory-Consciousness/dp/046506910X.Google Scholar
Ekman, P. (1999) Basic emotions. In: Handbook of cognition and emotion, ed. Dalgleish, T. & Powers, M. J., pp. 4560, John Wiley.CrossRefGoogle Scholar
Ekman, P. & Cordaro, D. T. (2011) What is meant by calling emotions basic. Emotion Review 3(4):364–70.CrossRefGoogle Scholar
Ellsworth, P. C. & Scherer, K. R. (2003) Appraisal processes in emotion. In: Handbook of affective sciences, ed. Davidson, R. J., Scherer, K. R. & Goldsmith, H., pp.572–95. Oxford University Press. Available at:http://www.amazon.com/Handbook-Affective-Sciences-Science/dp/0195126017.Google Scholar
Eslinger, P. J. & Damasio, A. R. (1985) Severe disturbance of higher cognition after bilateral frontal lobe ablation: Patient EVR. Neurology 35(12):1731–41. Available at:http://www.neurology.org/cgi/content/abstract/.CrossRefGoogle ScholarPubMed
Etkin, A. & Wager, T. D. (2007) Functional neuroimaging of anxiety: A meta-analysis of emotional processing in PTSD, social anxiety disorder, and specific phobia. The American Journal of Psychiatry 164:1476–88. Available at:http://ajp.psychiatryonline.org/cgi/content/abstract/164/10/1476.CrossRefGoogle ScholarPubMed
Ewbank, M. P., Barnard, P. J., Croucher, C. J., Ramponi, C. & Calder, A. J. (2009) The amygdala response to images with impact. Social Cognitive and Affective Neuroscience 4:127–33.CrossRefGoogle ScholarPubMed
Fanselow, M. S. & Poulos, A. M. (2005) The neuroscience of mammalian associative learning. Annual Review of Psychology 56:207–34. Available at:http://www.ncbi.nlm.nih.gov/pubmed/15709934 CrossRefGoogle ScholarPubMed
Feinstein, J. S., Adolphs, R., Damasio, A. & Tranel, D. (2011) The human amygdala and the induction and experience of fear. Current Biology 21:3438. Available at:http://www.ncbi.nlm.nih.gov/pubmed/21167712.CrossRefGoogle ScholarPubMed
Fendt, M. & Fanselow, M. S. (1999) The neuroanatomical and neurochemical basis of conditioned fear. Neuroscience and Biobehavioral Reviews 23:743–60. Available at:http://psycnet.apa.org/index.cfm?fa=search.displayRecord&uid=1999-05780-008.CrossRefGoogle ScholarPubMed
Ferris, C. F., Stolberg, T., Kulkarni, P., Murugavel, M., Blanchard, R., Blanchard, D. C., Febo, M., Brevard, M. & Simon, N. G. (2008) Imaging the neural circuitry and chemical control of aggressive motivation. BMC Neuroscience 9:135. Available at:http://www.biomedcentral.com/1471-2202/9/111.CrossRefGoogle ScholarPubMed
Ferry, A., Hespos, S. J. & Waxman, S. (2010) Language facilitates category formation in 3-month-old infants. Child Development 81:472–79. Available at:http://groups.psych.northwestern.edu/infantcognitionlab/FerryHesposWaxman2010.pdf.CrossRefGoogle Scholar
Fischer, H., Wright, C. I., Whalen, P. J., McInerney, S. C., Shin, L. M. & Rauch, S. L. (2003) Brain habituation during repeated exposure to fearful and neutral faces: A functional MRI study. Brain Research Bulletin 59:387–92. Available at:http://www.ncbi.nlm.nih.gov/pubmed/12507690.CrossRefGoogle ScholarPubMed
Fontaine, J. R. J., Scherer, K. R., Roesch, E. B. & Ellsworth, P. C. (2007) The world of emotions is not two-dimensional. Psychological Science 18:1050–57. Available at:http://pss.sagepub.com/content/18/12/1050.short.CrossRefGoogle Scholar
Fountoulakis, K. N., Giannakopoulos, P., Kövari, E. & Bouras, C. (2008) Assessing the role of cingulate cortex in bipolar disorder: Neuropathological, structural and functional imaging data. Brain Research Reviews 59:921.CrossRefGoogle ScholarPubMed
Fox, N. A. (1991) If it's not left, it's right: Electroencephalograph asymmetry and the development of emotion. The American Psychologist 46:863–72.CrossRefGoogle Scholar
Freedman, D. J., Riesenhuber, M., Poggio, T. & Miller, E. K. (2001) Categorical representation of visual stimuli in the primate prefrontal cortex. Science 291:312–16. Available at:http://www.sciencemag.org/content/291/5502/312.short.CrossRefGoogle ScholarPubMed
Freedman, D. J., Riesenhuber, M., Poggio, T. & Miller, E. K. (2002) Visual categorization and the primate prefrontal cortex: Neurophysiology and behavior. Journal of Neurophysiology 88:929. Available at:http://jn.physiology.org/cgi/content/abstract/88/2/929.CrossRefGoogle ScholarPubMed
Friston, L. (2010) The free energy principle: A unified brain theory? Nature Reviews Neuroscience 11:127–38.CrossRefGoogle ScholarPubMed
Fugate, J. M. B., Gouzoules, H. & Barrett, L. F. (2010) Reading chimpanzee faces: Evidence for the role of verbal labels in the categorical perception of emotion. Emotion 10:544–54. http://www.affective-science.org/pubs/2010/Fugate_Gouzoules_Barrett_2010CP.pdf.CrossRefGoogle ScholarPubMed
Fugate, J., Gouzoules, H. & Barrett, L. F. (2010) Reading chimpanzee faces: Evidence for the role of verbal labels in categorical perception of emotion. Emotion 10:544–54.CrossRefGoogle ScholarPubMed
Fusar-Poli, P., Placentino, A., Carletti, F., Landi, P., Allen, P., Surguladze, S., Benedetti, F., Abbamonte, M., Gasparotti, R., Barale, F., Perez, J., McGuire, P. & Politi, P. (2009) Functional atlas of emotional faces processing: A voxel-based meta-analysis of 105 functional magnetic resonance imaging studies. Journal of Psychiatry and Neuroscience 34(6):418–32. Available at:http://www.cma.ca/multimedia/staticContent/HTML/N0/l2/jpn/vol-34/issue-6/pdf/pg418.pdf.Google ScholarPubMed
Fuster, J. M. (2006) The cognit: A network model of cortical representation. International Journal of Psychophysiology 60:125–32. Available at:http://www.ncbi.nlm.nih.gov/pubmed/16626831.CrossRefGoogle ScholarPubMed
Gendron, M. & Barrett, L. F. (2009) Reconstructing the past: A century of ideas about emotion in psychology. Emotion Review 1(4):316–39. Available at:http://www2.bc.edu/~barretli/pubs/2009/gendron-barrett-2009.pdf.CrossRefGoogle Scholar
Gendron, M., Lindquist, K. A., Wager, T. D., Bliss-Moreau, E., Kober, H. & Barrett, L. F. (in preparation) Seeing with feeling: A meta-analytic review of visual cortex activity in emotion.Google Scholar
Gitelman, D. R., Nobre, A. C., Sonty, S., Parrish, T. B. & Mesulam, M. M. (2005) Language network specializations: An analysis with parallel task designs and functional magnetic resonance imaging. Neuroimage 26:975–85. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/15893473.CrossRefGoogle ScholarPubMed
Glenn, A. L., Raine, A. & Schug, R. A. (2009) The neural correlates of moral decision making in psychopathy. Molecular Psychiatry 14:59. Available at:http://www.nature.com/mp/journal/v14/n1/abs/mp2008104a.html.CrossRefGoogle ScholarPubMed
Gloor, P. (1990) Experiential phenomena of temporal lobe epilepsy. Brain 113:1673–94.CrossRefGoogle ScholarPubMed
Gotlib, I. H. & Hamilton, J. P. (2008) Neuroimaging and depression: Current status and unresolved issues. Current Directions in Psychological Science 17:159. Available at:http://psycnet.apa.org/?fa=main.doiLanding&uid=2008-04591-019.CrossRefGoogle Scholar
Goyer, P. F., Konicki, P. E. & Schulz, S. C. (1994) Brain imaging in personality disorders. In: Biological and neurobehavioral studies of borderline personality disorder, ed. Silk, K. M., pp. 109–25.Google Scholar
Grafman, J., Schwab, K., Warden, D., Pridgen, A., Brown, H. R. & Salazar, A. M. (1996) Frontal lobe injuries and violence a report of the Vietnam Head Injury Study. Neurology 46:1231–38. Available at:http://www.ncbi.nlm.nih.gov/pubmed/8628458.CrossRefGoogle ScholarPubMed
Greene, J. D., Nystrom, L. E., Engell, A. D., Darley, J. M. & Cohen, J. D. (2004) The neural bases of cognitive conflict and control in moral judgment. Neuron 44:389400. Available at:http://www.wjh.harvard.edu/~jgreene/GreeneWJH/Greene-etal-Neuron04.pdf.CrossRefGoogle ScholarPubMed
Gregg, T. R. & Siegel, A. (2001) Brain structures and neurotransmitters regulating aggression in cats: Implications for human aggression. Progress in Neuropsychopharacology Biological Psychiatry 25:91140. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/11263761.CrossRefGoogle ScholarPubMed
Grezes, J. & Decety, J. (2001) Functional anatomy of execution, mental simulation, observation, and verb generation of actions: A meta-analysis. Human Brain Mapping 12:119. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/11198101.3.0.CO;2-V>CrossRefGoogle ScholarPubMed
Grinband, J., Savitsky, J., Wager, T. D., Teichert, T., Ferrera, V. P. & Hirsch, J. (2011) The dorsal medial frontal cortex is sensitive to time on task, not response conflict or error likelihood. Neuroimage 57(2):303–11.CrossRefGoogle ScholarPubMed
Gross, J. J. & Barrett, L. F. (2011) Emotion generation and emotion regulation: One or two depends on your point of view. Emotion Review 3:816.CrossRefGoogle ScholarPubMed
Halgren, E., Walter, R. D., Cherlow, D. G. & Crandall, P. H. (1978) Mental phenomena evoked by electrical stimulation of the human hippocampal formation and amygdala. Brain 101:83117. Available at:http://www.ncbi.nlm.nih.gov/pubmed/638728.CrossRefGoogle ScholarPubMed
Harenski, C. L., Kim, S. H. & Hamann, S. (2009) Neuroticism and psychopathy predict brain activation during moral and nonmoral emotion regulation. Cognitive, Affective, and Behavioral Neuroscience 9(1):115. Available at:http://www.ncbi.nlm.nih.gov/pubmed/19246323.CrossRefGoogle ScholarPubMed
Harlow, H. F. & Stagner, R. (1932) Psychology of feelings and emotions. I. Theory of feelings. Psychological Review 39:570–89. Available at:http://psycnet.apa.org/journals/rev/39/6/570.pdf.CrossRefGoogle Scholar
Harmon-Jones, E. & Allen, J. J. (1998) Anger and frontal brain activity: EEG asymmetry consistent with approach motivation despite negative affective valence. Journal of Personality and Social Psychology 74:1310–16. Available at:http://psycnet.apa.org/index.cfm?fa=fulltext.journal&jcode=psp&vol=74&issue=5&format=html&page=1310&expand=1.CrossRefGoogle ScholarPubMed
Harmon-Jones, E. & Sigelman, J. (2001) State anger and prefrontal brain activity: Evidence that insult-related relative left-prefrontal activation is associated with experienced anger and aggression. Journal of Personality and Social Psychology 80:797803. Available at:http://psycnet.apa.org/journals/psp/80/5/797.html.CrossRefGoogle ScholarPubMed
Harrison, N. A., Gray, M. A., Giarnos, P. J. & Critchley, H. G. (2010) The embodiment of emotional feelings in the brain. The Journal of Neuroscience 30:12878–84. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/20861391.CrossRefGoogle ScholarPubMed
Hassabis, D. & Maguire, E. A. (2009) The construction system of the brain. Philosophical Transactions of the Royal Society B 364:1263–71. Available at:http://rstb.royalsocietypublishing.org/content/364/1521/1263.full.CrossRefGoogle ScholarPubMed
Haxby, J. V., Gobbini, M. I., Furey, M. L., Ishai, A., Schouten, J. L. & Pietrini, P. (2001) Distributed and overlapping representations of faces and objects in ventral temporal cortex. Science 293:2425–30. Available at:http://www.sciencemag.org/cgi/content/abstract/sci;293/5539/2425.CrossRefGoogle ScholarPubMed
Herry, C., Bach, D. R., Esposito, F., Di Salle, F., Perrig, W. J., Scheffler, K., Lüthi, A. & Seifritz, E. (2007) Processing of temporal unpredictability in human and animal amygdala. Journal of Neuroscience 27:5958. Available at:http://www.ncbi.nlm.nih.gov/pubmed/17537966.CrossRefGoogle ScholarPubMed
Hitchcock, J. M. & Davis, M. (1986) Lesions of the amygdala, but not of the cerebellum or red nucleus, block conditioned fear as measured with the potentiated startle paradigm. Behavioral Neuroscience 100:1122. Available at:http://psycnet.apa.org/journals/bne/100/1/11.html.CrossRefGoogle ScholarPubMed
Hitchcock, J. M. & Davis, M. (1987) Fear-potentiated startle using an auditory conditioned stimulus: Effect of lesions of the amygdala. Physiology and Behavior 39:403408. Available at:http://www.ncbi.nlm.nih.gov/pubmed/3575483.CrossRefGoogle ScholarPubMed
Holland, P. C. & Gallagher, M. (1999) Amygdala circuitry in attentional and representational processes. Trends in Cognitive Sciences 3:6573. Available at:http://www.ncbi.nlm.nih.gov/pubmed/10234229.CrossRefGoogle ScholarPubMed
Hornak, J., Bramham, J., Rolls, E. T., Morris, R. G., O'Doherty, J., Bullock, P. R. & Polkey, C. E. (2003) Changes in emotion after circumscribed surgical lesions of the orbitofrontal and cingulate cortices. Brain: A Journal of Neurology 126:1691–712. Available at:http://www.ncbi.nlm.nih.gov/pubmed/12805109.CrossRefGoogle ScholarPubMed
Hornak, J., O'Doherty, J., Bramham, J., Rolls, E. T., Morris, R. G., Bullock, P. R. & Polkey, C. E. (2004) Reward-related reversal learning after surgical excisions in orbito-frontal or dorsolateral prefrontal cortex in humans. Journal of Cognitive Neuroscience 16:463–78.CrossRefGoogle ScholarPubMed
Hunt, W. A. (1941) Recent developments in the field of emotion. Psychological Bulletin 38:249–76. Available at:http://psycnet.apa.org/?&fa=main.doiLanding&doi=10.1037/h0054615.CrossRefGoogle Scholar
Izard, C. E. (2011) Forms and functions of emotions: Matters of emotion-cognition interactions. Emotion Review 3:371–78.CrossRefGoogle Scholar
Jabbi, M., Bastiaansen, J. & Keysers, C. (2008) A common anterior insula representation of disgust observation, experience and imagination shows divergent functional connectivity pathways. PLoS One 3:18.CrossRefGoogle ScholarPubMed
James, W. (1884) What is an emotion? Mind 9:188205. Available at:http://www.jstor.org/pss/2246769.CrossRefGoogle Scholar
James, W. (1890/;1998) Principles of psychology. University of Chicago Press.Google Scholar
Jenison, R. L., Rangel, A., Oya, H., Kawasaki, H. & Howard, M. A. (2011) Value encoding in single neurons in the human amygdala during decision making. The Journal of Neuroscience 31:331–38. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/21209219.CrossRefGoogle ScholarPubMed
Jürgens, U. (2009) The neural control of vocalization in mammals: A review. Journal of Voice 23:110. Available at:http://www.ncbi.nlm.nih.gov/pubmed/8653176.CrossRefGoogle ScholarPubMed
Kamback, M. (1973) The effects of orbital and dorsolateral frontal cortical ablations on ethanol self-selection and emotional behaviors in monkeys. Neuropsychologia 11:331–35. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/4209334.CrossRefGoogle ScholarPubMed
Keltner, D., Ekman, P., Gonzaga, G. & Beer, J. (2003) Facial expressions of emotion. In: Handbook of affective sciences, ed. Davidson, R. J., Scherer, K. R. & Goldsmith, H. H., pp. 415–27. Oxford University Press. Available at:http://www.amazon.com/Handbook-Affective-Sciences-Science/dp/0195126017.Google Scholar
Kennedy, D. P. & Adolphs, R. (2010) Impaired fixation to eyes following amygdala damage arises from abnormal bottom-up attention. Neuropsychologia 48:3392–98. Available at:http://www.ncbi.nlm.nih.gov/pubmed/20600184.CrossRefGoogle ScholarPubMed
Kensinger, E. A. & Corkin, S. (2004) Two routes to emotional memory: Distinct neural processes for valence and arousal. Proceedings of the National Academy of Sciences USA 101:3310–15. Available at:http://www2.bc.edu/elizabeth-kensinger/Kensinger_Corkin_PNAS04.pdf.CrossRefGoogle ScholarPubMed
Kim, J. J., Rison, R. A. & Fanselow, M. S. (1993) Effects of amygdala, hippocampus, and periaqueductal gray lesions on short- and long-term contextual fear. Behavioral Neuroscience 107:1093–98. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/8136063.CrossRefGoogle Scholar
Kipps, C. M., Duggins, A. J., McCusker, E. A. & Calder, A. J. (2007) Disgust and happiness recognition correlate with anteroventral insula and amygdala volume respectively in pre-clinical Huntington's disease. Journal of Cognitive Neuroscience 19:1206–217. Available at:http://www.ncbi.nlm.nih.gov/pubmed/17583995.CrossRefGoogle Scholar
Knutson, B., Rick, S., Wimmer, G. E., Prelec, D. & Loewenstein, G. (2007) Neural predictors of purchases. Neuron 53:147–56. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1876732/.CrossRefGoogle ScholarPubMed
Kober, H., Barrett, L. F., Joseph, J., Bliss-Moreau, E., Lindquist, K. & Wager, T. D. (2008) Functional grouping and cortical-subcortical interactions in emotion: A meta-analysis of neuroimaging studies. NeuroImage 42:9981031. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2752702/.CrossRefGoogle ScholarPubMed
Koenigs, M., Young, L., Adolphs, R., Tranel, D., Cushman, F., Hauser, M. & Damasio, A. (2007) Damage to the prefrontal cortex increases utilitarian moral judgments. Nature 446:908–11. Available at:http://www.ncbi.nlm.nih.gov/pubmed/17377536.CrossRefGoogle Scholar
Kopchia, K. L., Altman, H. J. & Commissaris, R. L. (1992) Effects of lesions of the central nucleus of the amygdala on anxiety-like behaviors in the rat. Pharmacology, Biochemistry, and Behavior 43:453–61. Available at:http://www.ncbi.nlm.nih.gov/pubmed/1438482.CrossRefGoogle ScholarPubMed
Krain, A. L., Wilson, A. M., Arbuckle, R., Castellanos, F. X. & Milham, M. P. (2006) Distinct neural mechanisms of risk and ambiguity: A meta-analysis of decision-making. Neuroimage 32:477–84. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/16632383.CrossRefGoogle ScholarPubMed
Kringelbach, M. L. & Rolls, E. T. (2004) The functional neuroanatomy of the human orbitofrontal cortex: Evidence from neuroimaging and neuropsychology. Progress in Neurobiology 72:341–72. Available at:http://www.kringelbach.dk/papers/PN_Kringelbach2004.pdf.CrossRefGoogle ScholarPubMed
LaBar, K. S., Gatenby, J. C., Gore, J. C., LeDoux, J. E. & Phelps, E. A. (1998) Human amygdala activation during conditioned fear acquisition and extinction: A mixed-trial fMRI study. Neuron 20:937–45. Available at:http://www.ncbi.nlm.nih.gov/pubmed/9620698.CrossRefGoogle ScholarPubMed
LaBar, K. S., LeDoux, J. E., Spencer, D. D. & Phelps, E. A. (1995) Impaired fear conditioning following unilateral temporal lobectomy in humans. Journal of Neuroscience 15(10):6846–55. Available at:http://www.ncbi.nlm.nih.gov/pubmed/7472442.CrossRefGoogle ScholarPubMed
Laine, C. M., Spitler, K. M., Mosher, C. P. & Gothard, K. M. (2009) Behavioral triggers of skin conductance responses and their neural correlates in the primate amygdala. Journal of Neurophysiology 101:1749–54. Available at:http://www.ncbi.nlm.nih.gov/pubmed/19144740.CrossRefGoogle ScholarPubMed
Lambon Ralph, M. A., Pobric, G. & Jefferies, E. (2009) Conceptual knowledge is underpinned by the temporal pole bilaterally: Convergent evidence from rTMS. Cerebral Cortex 19:832–38. Available at:http://www.ncbi.nlm.nih.gov/pubmed/18678765.CrossRefGoogle ScholarPubMed
Lang, P. J., Davis, M. & Öhman, A. (2000) Fear and anxiety: Animal models and human cognitive psychophysiology. Journal of Affective Disorders 61:137–59. Available at:http://www.stanford.edu/~kateri/Becky/PDFs/Lang%202000.pdf.CrossRefGoogle ScholarPubMed
LeDoux, J. (2007) The amygdala. Current Biology 17:R868–74.CrossRefGoogle ScholarPubMed
LeDoux, J. E., Ciccetti, P., Xagoraris, A. & Romanski, L. M. (1990) The lateral amygdaloid nucleus: Sensory interface of the amygdala in fear conditioning. Journal of Neuroscience 10:1062–69. Available at:http://www.ncbi.nlm.nih.gov/pubmed/2329367.CrossRefGoogle ScholarPubMed
LeDoux, J. E., Sakaguchi, A., Iwata, J. & Reis, D. J. (1985) Auditory emotional memories: Establishment by projections from the medial geniculate nucleus to the posterior neostriatum and/or dorsal amygdala. Annals of the New York Academy of Sciences 444:463–64. Available at:http://www.ncbi.nlm.nih.gov/pubmed/3860099.CrossRefGoogle ScholarPubMed
LeDoux, J. E., Sakaguchi, A. & Reis, D.J (1983) Strain differences in fear between spontaneously hypertensive and normotensive rats. Brain Research 277:137–43. Available at:http://www.ncbi.nlm.nih.gov/pubmed/6640286.CrossRefGoogle ScholarPubMed
Lindquist, K. A. & Barrett, L. F. (2008a) Constructing emotion: The experience of fear as a conceptual act. Psychological Science 19:898903. Available at:http://www2.bc.edu/~lindqukr/docs/Lindquist&Barrett2008.pdf.CrossRefGoogle ScholarPubMed
Lindquist, K. A., Barrett, L. F., Bliss-Moreau, E. & Russell, J. A. (2006) Language and the perception of emotion. Emotion 6:125–38. Available at:http://nmr.mgh.harvard.edu/~lindqukr/publications.html.CrossRefGoogle ScholarPubMed
Lovick, T. A. (1992) Inhibitory modulation of the cardiovascular defence response by the ventrolateral periaqueductal grey matter in rats. Experimental Brain Research 89:133–39. Available at:http://www.springerlink.com/content/h18327q02g311765/.CrossRefGoogle ScholarPubMed
Machado, C. J. & Bachevalier, J. (2006) The impact of selective amygdala, orbital frontal cortex, or hippocampal formation lesions on established social relationships in rhesus monkeys (Macaca mulatta). Behavioral Neuroscience 120:761–86. Available at:http://www.ncbi.nlm.nih.gov/pubmed/16893284.CrossRefGoogle ScholarPubMed
MacLean, P. D. & Newman, J. D. (1988) Role of midline frontolimbic cortex in production of the isolation call of squirrel monkeys. Brain Research 450:111–23. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/3401705.CrossRefGoogle ScholarPubMed
Mandler, G. (1975) Mind and emotion. Wiley.Google Scholar
Mandler, G. (1990) William James and the construction of emotion. Psychological Science 1:179–80. Available at:http://www.jstor.org/stable/40062633.CrossRefGoogle Scholar
Mansouri, F. A., Tanaka, K. & Buckley, M. J. (2009) Conflict-induced behavioral adjustment: A clue to the executive functions of prefrontal cortex. Nature Reviews Neuroscience 10:141–52. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/19153577.CrossRefGoogle Scholar
Mantyh, P. W. (1983) Connections of midbrain periaqueductal gray in the monkey. I. Ascending efferent projections. Journal of Neurophysiology 49(3):567–81. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/6300350.CrossRefGoogle ScholarPubMed
Maskati, A. A. A. & Zbrozyna, A. W. (1989) Cardiovascular and motor components of the defense reaction elicited in rats by electrical and chemical stimulation in amygdala. Journal of the Autonomic Nervous System 28:127–32. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/2625501.CrossRefGoogle ScholarPubMed
Mason, W. A., Capitanio, J. P., Machado, C. J., Mendoza, S. P. & Amaral, D. G. (2006) Amygdalectomy and responsiveness to novelty in rhesus monkeys (Macaca mulatta): Generality and individual consistency of effects. Emotion 6:7381.CrossRefGoogle ScholarPubMed
Mauss, I. B. & Robinson, M. D. (2009) Measures of emotion: A review. Cognition and Emotion 23:209–37. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2756702/.CrossRefGoogle ScholarPubMed
Mayberg, H. S., Lozano, A. M., Voon, V., McNeely, H. E., Seminowicz, D., Hamani, C., Schwalb, D. & Kennedy, S. H. (2005) Deep brain stimulation for treatment-resistant depression. Neuron 45:651–60. Available at:http://www.ncbi.nlm.nih.gov/pubmed/15748841.CrossRefGoogle ScholarPubMed
McDermott, K. B., Szpunar, K. K. & Christ, S. E. (2009) Laboratory-based and autobiographical retrieval tasks differ substantially in their neural substrates. Neuropsychologia 47:2290–98. Available at:http://memory.wustl.edu/McDermott_Lab/Publications.html.CrossRefGoogle ScholarPubMed
McIntosh, A. R. (2004) Contexts and catalysts: A resolution of the location and integration of function in the brain. Neuroinformatics 2:175–81 Available at:http://www.springerlink.com.ezp-prod1.hul.harvard.edu/content/k12p31883927q237/.CrossRefGoogle Scholar
McIntosh, A. R., Bookstein, F. L., Haxby, J. V. & Grady, C. L. (1996) Spatial pattern analysis of functional brain images using partial least squares. Neuroimage 3:143–57. Available at:http://www.ncbi.nlm.nih.gov/pubmed/9345485.CrossRefGoogle ScholarPubMed
Mendes, W. B., Blascovich, J., Hunter, S. B., Lickel, B. & Jost, J. T. (2007) Threatened by the unexpected: Physiological responses during social interactions with expectancy-violating partners. Journal of Personality and Social Psychology 92:698. Available at:http://psycnet.apa.org/journals/psp/92/4/698.html.CrossRefGoogle ScholarPubMed
Mesulam, M. M. & Mufson, E. J. (1982) Insula of the old world monkey. 111: Efferent cortical output and comments on function. The Journal of Comparative Neurology 212:3852. Available at:http://www3.interscience.wiley.com/journal/109687236/abstract.CrossRefGoogle Scholar
Milders, M., Crawford, J. R., Lamb, A. & Simpson, S. A. (2003) Differential deficits in expression recognition in gene-carriers and patients with Huntington's disease. Neuropsychologia 41:1484–92. Available at:http://www.ncbi.nlm.nih.gov/pubmed/12849766.CrossRefGoogle ScholarPubMed
Milham, M. P., Banich, M. T., Webb, A., Barad, V., Cohen, N. J., Wszalek, T. & Kramer, A. F. (2001) The relative involvement of anterior cingulate and prefrontal cortex in attentional control depends on nature of conflict. Brain Research. Cognitive Brain Research 12:467–73. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/11689307.CrossRefGoogle ScholarPubMed
Miller, E. K. (2000) The prefrontal cortex and cognitive control. Nature Reviews Neuroscience 1:5965. Available at:http://www.doc.ic.ac.uk/~xh1/Referece/Visual-search/neural-cognitive-contrl.pdf.CrossRefGoogle ScholarPubMed
Miller, E. K. & Cohen, J. D. (2001) An integrative theory of prefrontal cortex function. Annual Review of Neuroscience 24:167202. Available at:http://arjournals.annualreviews.org/doi/abs/10.1146%2Fannurev.neuro.24.1.167.CrossRefGoogle ScholarPubMed
Miller, E. K., Freedman, D. J. & Wallis, J. D. (2002) The prefrontal cortex: Categories, concepts and cognition. Philosophical Transactions: Biological Sciences 357:1123–36. Available at:http://rstb.royalsocietypublishing.org/content/357/1424/1123.full.pdf.CrossRefGoogle ScholarPubMed
Missilin, R. & Ropartz, P. (1981) Effects of lateral amygdala lesions on the responses to novelty in mice. Behavioural Processes 6:329–36.CrossRefGoogle Scholar
Mitchell, I. J., Heims, H., Neville, E. A. & Rickards, H. (2005) Huntington's disease patients show impaired perception of disgust in the gustatory and olfactory modalities. Journal of Neuropsychiatry and Clinical Neurosciences 17:119. Available at:http://www.ncbi.nlm.nih.gov/pubmed/15746492.CrossRefGoogle ScholarPubMed
Mitchell, J. P. (2009) Inferences about other people's minds. Philosophical Transactions of the Royal Society B 364:1309–16. Available at:http://www.wjh.harvard.edu/~scanlab/papers/2009_predictions_PhilTrans.pdf.CrossRefGoogle Scholar
Mobbs, D., Petrovic, P., Marchant, J. L., Hassabis, D., Weiskopf, N., Seymour, B., Dolan, R. J. & Frith, C. D. (2007) When fear is near: Threat imminence elicits prefrontal periaqueductal gray shifts in humans. Science 317(5841):1079–83. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2648508/.CrossRefGoogle ScholarPubMed
Moriguchi, Y., Negreira, A., Weirerich, M., Dautoff, R., Dickerson, B. C., Wright, C. I. & Barrett, L. F. (2010) Differential hemodynamic response in affective circuitry with aging: Novely, valence and arousal. Journal of Cognitive Neuroscience 23:1027–41. Available at:http://www.mitpressjournals.org.ezp-prod1.hul.harvard.edu/doi/pdf/10.1162/jocn.2010.21527.Google Scholar
Moulton, S. T. & Kosslyn, S. (2009) Imagining predictions: Mental imagery as mental emulation. Philosophical Transactions of the Royal Society B 364:1273–80. Available at:http://rstb.royalsocietypublishing.org/content/364/1521/1273.full.CrossRefGoogle ScholarPubMed
Moyer, K. E. (1968) Kinds of aggression and their physiological basis. Communications in Behavioral Biology A2: 6587.Google Scholar
Murphy, F. C., Nimmo-Smith, I. & Lawrence, A. D. (2003) Functional neuroanatomy of emotions: A meta-analysis. Cognitive, Affective, and Behavioral Neuroscience, 3(3):207–33.CrossRefGoogle ScholarPubMed
Murray, E. A. (2007) The amygdala, reward and emotion. Trends in Cognitive Science 11:489–97.CrossRefGoogle ScholarPubMed
Nachman, M. & Ashe, J. H. (1974) Effects of basolateral amygdala lesions on neophobia, learned taste aversions, and sodium appetite in rats. Journal of Comparative and Physiological Psychology 87:622–43. Available at:http://psycnet.apa.org/journals/com/87/4/622.pdf.CrossRefGoogle ScholarPubMed
Nelson, J. K., Reuter-Lorenz, P. A., Sylvester, C. C., Jonides, J. & Smith, E. E. (2003) Dissociable neural mechanisms underlying response-based and familiarity-based conflict in working memory. Proceedings of the National Academy of Sciences USA 100:11171–75. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/12958206.CrossRefGoogle ScholarPubMed
Nelson, S. M., Dosenbach, N. U. F., Cohen, A. L., Wheeler, M. E., Schlaggar, B. L. & Petersen, S. E. (2010) Role of the anterior insula in task-level control and focal attention. Brain Structure and Function 214:669–80.CrossRefGoogle ScholarPubMed
Newman, S. W. (1999) The medial extended amygdala in male reproductive behavior. A node in the mammalian social behavior network. Annals of the New York Academy of Sciences 877:242–57. Available at:http://www.ncbi.nlm.nih.gov/sites/entrez.CrossRefGoogle ScholarPubMed
Northoff, (2006) Is emotion regulation self-regulation? Trends in Cognitive Sciences 9:408409. Available at:http://dept.psych.columbia.edu/~kochsner/pdf/Northoff_Tics_comment.pdf.CrossRefGoogle Scholar
Numan, M. (2007) Motivational systems and the neural circuitry of maternal behavior in the rat. Developmental Psychobiology 49:12–21. Available at:http://www.ncbi.nlm.nih.gov/pubmed/17186513.CrossRefGoogle ScholarPubMed
Ochsner, K. N., Knierim, K., Ludlow, D., Hanelin, J., Ramachandran, T. & Mackey, S. (2004a) Reflecting upon feelings: An fMRI study of neural systems supporting the attribution of emotion to self and other. Journal of Cognitive Neuroscience 16(10):1748–72. Available at:http://dept.psych.columbia.edu/~kochsner/pdf/Ochsner_Reflect_Feelings.pdf.CrossRefGoogle ScholarPubMed
Ochsner, K. N., Ray, R. D., Cooper, J. C., Robertson, E. R., Chopra, S., Gabrieli, J. D. & Gross, J. J. (2004b) For better or for worse: neural systems supporting the cognitive down- and up-regulation of negative emotion. NeuroImage 23(2):483–99. Available at:http://www.ncbi.nlm.nih.gov/pubmed/15488398.CrossRefGoogle ScholarPubMed
Öhman, A. (2009) Of snakes and faces: An evolutionary perspective on the psychology of fear. Scandinavian Journal of Psychology 50:543–52. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/19930253.CrossRefGoogle ScholarPubMed
Öngür, D. & Price, J. L. (2000) The organization of networks within the orbital and medial prefrontal cortex of rats, monkeys and humans. Cerebral Cortex 10(3):206–19. Available at:http://cercor.oxfordjournals.org/cgi/content/abstract/10/3/206.CrossRefGoogle ScholarPubMed
Ortigue, S., Grafton, S. T. & Bianchi-Demicheli, F. (2007) Correlation between insula activation and self-reported quality of orgasm in women. NeuroImage 37:551–60. Available at:http://www.ncbi.nlm.nih.gov/pubmed/17601749.CrossRefGoogle ScholarPubMed
Ortony, A. & Turner, T. J. (1990) What's basic about basic emotions? Psychological Review 97:315–31.CrossRefGoogle ScholarPubMed
Owen, A. M., McMillan, K., Laird, A. R. & Bullmore, E. (2005) N-Back working memory paradigm: A meta-analysis of normative functional neuroimaging studies. Human Brain Mapping 25:4659. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/15846822.CrossRefGoogle ScholarPubMed
Panksepp, J. (1998) Affective neuroscience: The foundations of human and animal emotions. Oxford University Press.CrossRefGoogle Scholar
Panksepp, J. (2007) Neurologizing the psychology of affects: How appraisal-based constructivism and basic emotion theory can coexist. Perspectives on Psychological Science 2(3):281–96. Available at:http://www3.interscience.wiley.com/journal/118509152/abstract.CrossRefGoogle ScholarPubMed
Panksepp, J. & Watt, D. (2011) What is basic about basic emotions? Lasting lessons from affective neuroscience. Emotion Review 3(4):387–96.CrossRefGoogle Scholar
Paton, J. P., Belova, M. A., Morrison, S. E. & Salzman, C. D. (2006) The primate amygdala represents the positive and negative value of visual stimuli during learning. Nature 16:865–70. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2396495/.CrossRefGoogle Scholar
Paus, T. (2001) Primate anterior cingulate cortex: Where motor control, drive and cognition interface. Nature Reviews Neuroscience 2:417–24. Available at:http://www.oise.utoronto.ca/research/brainwaves/phpwebsite/files/uplink/Paus_2001.pdf.CrossRefGoogle ScholarPubMed
Penfield, W. & Falk, M. E. (1955) The insula. Further observations on its function. Brain 78:445–70. Available at:http://brain.oxfordjournals.org/content/78/4/445.extract.CrossRefGoogle ScholarPubMed
Pessoa, L. (2008) On the relationship between emotion and cognition. Nature Reviews Neuroscience 9(2):148–58.CrossRefGoogle ScholarPubMed
Pessoa, L. (2010b) Emotion and cognition and the amygdala: From “what is it?” to “what's to be done?”. Neuropsychologia 48:3416–29. Available at:http://www.indiana.edu/%7Elceiub/publications_files/Pessoa_Neuropsychologia_2010.pdf.CrossRefGoogle Scholar
Pessoa, L. & Adolphs, R. (2010) Emotion processing and the amygdala: From a “low road” to “many roads” of evaluating biological significance. Nature Reviews Neuroscience 11:773–83.CrossRefGoogle Scholar
Petrides, M. (2005) Lateral prefrontal cortex: Architectonic and functional organization. Philosophical Transactions of the Royal Society of London, B: Biological Sciences 360:781–95. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/15937012.CrossRefGoogle ScholarPubMed
Petrides, M. (2007) The orbitofrontal cortex: Novelty, deviation from expectation, and memory. Annals of the New York Academy of Sciences 1121:3353. Available at:http://www.ncbi.nlm.nih.gov/pubmed/17872393.CrossRefGoogle ScholarPubMed
Phan, K. L., Wager, T., Taylor, S. F. & Liberzon, I. (2002) Functional neuroanatomy of emotion: A meta-analysis of emotion activation studies in PET and fMRI. NeuroImage 16(2):331–48. Available at:http://www.ncbi.nlm.nih.gov/pubmed/12030820.CrossRefGoogle ScholarPubMed
Phelps, E. A., Ling, S. & Carrasco, M. (2006). Emotion facilitates perception and potentiates the perceptual benefits of attention. Psychological Science 17(4):292–99.CrossRefGoogle ScholarPubMed
Pobric, G., Jefferies, E. & Ralph, M. A. L. (2007) Anterior temporal lobes mediate semantic representation: Mimicking semantic dementia by using rTMS in normal participants. Proceedings of the National Academy of Science 104:20137–41. Available at:http://www.pnas.org/content/104/50/20137.abstract.CrossRefGoogle ScholarPubMed
Poldrack, R. A., Halchenko, Y. O. & Hanson, S. J. (2009) Decoding the large-scale structure of brain function by classifying mental States across individuals. Psychological Science 20:1364–72. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/19883493.CrossRefGoogle ScholarPubMed
Price, C. J. & Friston, K. J. (2005) Functional ontologies for cognition: The systematic definition of structure and function. Cognitive Neuropsychology 22:262–75. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/21038249.CrossRefGoogle ScholarPubMed
Raichle, M. E., MacLeod, A. M., Snyder, A. Z., Powers, W. J., Gusnard, D. A. & Shulman, G. L. (2001) A default mode of brain function. Proceedings of the National Academy of Sciences USA 98:676–82. Available at:http://www.pnas.org/content/98/2/676.long.CrossRefGoogle ScholarPubMed
Raine, A., Lencz, T., Bihrle, S., LaCasse, L. & Colletti, P. (2000) Reduced prefrontal gray matter volume and reduced autonomic activity in antisocial personality disorder. Archives of General Psychiatry 57:119–27. Available at:http://www.ncbi.nlm.nih.gov/pubmed/10665614.CrossRefGoogle ScholarPubMed
Raleigh, M. J., Steklis, H. D., Ervin, F. R., Kling, A. S. & McGuire, M. T. (1979) The effects of orbitofrontal lesions on the aggressive behavior of vervet monkeys (Cercopithecus aethiops sabaeus). Experimental Neurology 66:158–68. Available at:http://www.ncbi.nlm.nih.gov/pubmed/113236.CrossRefGoogle ScholarPubMed
Rankin, K. P., Gorno-Tempini, M. L., Allison, S. C., Stanley, C. M., Glenn, S., Weiner, M. W. & Miller, B. L. (2006) Structural anatomy of empathy in neurodegenerative disease. Brain 129:2945–56.CrossRefGoogle ScholarPubMed
Rempel-Clower, N. L. & Barbas, H. (1998) Topographic organization of connections between the hypothalamus and prefrontal cortex in the rhesus monkey. Journal of Comparative Neurology 398:393419. Available at:http://www.ncbi.nlm.nih.gov/pubmed/9714151.3.0.CO;2-V>CrossRefGoogle ScholarPubMed
Reynolds, S. M. & Berridge, K. C. (2002) Positive and negative motivation in nucleus accumbens shell: Bivalent rostrocaudal gradients for GABA-elicited eating, taste “liking”/”disliking” reactions, place preference/avoidance, and fear. The Journal of Neuroscience 22:7308–20. Available at:http://www.jneurosci.org/cgi/content/abstract/22/16/7308.CrossRefGoogle Scholar
Reynolds, S. M. & Berridge, K. C. (2003) Glutamate motivational ensembles in nucleus accumbens: Rostrocaudal shell gradients of fear and feeding. European Journal of Neuroscience 17:2187–200. Available at:http://www.ncbi.nlm.nih.gov/pubmed/12786986.CrossRefGoogle ScholarPubMed
Reynolds, S. M. & Berridge, K. C. (2008) Emotional environments retune the valence of appetitive versus fearful functions in nucleus accumbens. Nature Neuroscience 11:423–25. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2717027/.CrossRefGoogle ScholarPubMed
Roberson, D., Damjanovic, L. & Pilling, M. (2007). Categoricalperception of expressions: Evidence for a “category adjustment” model. Memory and Cognition 35:1814–29.CrossRefGoogle Scholar
Roberson, D. & Davidoff, J. (2000) The categorical perception of color and facial expressions: The effect of verbal interference. Memory and Cognition 28:977–86. Available at:http://www.ncbi.nlm.nih.gov/pubmed/11105523.CrossRefGoogle ScholarPubMed
Rogers, T. T., Lambon Ralph, M. A., Garrard, P., Bozeat, S., McClelland, J. L., Hodges, J. R. & Patterson, K. (2004) Structure and deterioration of semantic memory: A neuropsychological and computational investigation. Psychological Review 111:205–35. Available at:http://charlotte.cnbc.cmu.edu/ibsc/papers/RogersETAL04_PR.pdf.CrossRefGoogle ScholarPubMed
Rolls, E. T. (1999) The function of the orbitofrontal cortex. Neurocase 5:301–12. Available at:http://www.oxcns.org/papers/357_Rolls04.pdf.CrossRefGoogle Scholar
Rolls, E. T., Critchley, H. D., Mason, R. & Wakeman, E. A. (1996) Orbitofrontal cortex neurons: Role in olfactory and visual association learning. Journal of Neurophysiology 75:1970–81. Available at:http://www.ncbi.nlm.nih.gov/pubmed/8734596.CrossRefGoogle ScholarPubMed
Rolls, E. T., Hornak, J., Wade, D. & McGrath, J. (1994) Emotion-related learning in patients with social and emotional changes associated with frontal lobe damage. Journal of Neurology, Neurosurgery and Psychiatry 57:1518–24. Available at:http://www.ncbi.nlm.nih.gov/pubmed/7798983.CrossRefGoogle ScholarPubMed
Roseman, I. J. (1984) Cognitive determinants of emotion: A structural theory. Review of Personality and Social Psychology 5:1136.Google Scholar
Rosen, H. J., Pace-Savitsky, K., Perry, R. J., Kramer, J. H., Miller, B. L. & Levenson, R. W. (2004) Recognition of emotion in the frontal and temporal variants of frontotemporal dementia. Dementia and Geriatric Cognitive Disorders 17:277–81. Available at:http://www.ncbi.nlm.nih.gov/pubmed/15178936.CrossRefGoogle ScholarPubMed
Rosen, J. B. & Davis, M. (1988) Enhancement of acoustic startle by electrical stimulation of the amygdala. Behavioral Neuroscience 102:195202.CrossRefGoogle ScholarPubMed
Rozin, P., Haidt, J. & McCaule, C. (2008) Disgust. In: The handbook of emotions, 3rd edition, ed. Lewis, M., Haviland-Jones, J. M. & Barrett, L. F., pp. 757–76. Guilford Press.Google Scholar
Rozin, P., Haidt, J. & McCauley, C. (2000) Disgust. In: Handbook of emotions, 2nd edition, ed. Lewis, M. & Havilland-Jones, J. M., pp. 637–53. Guilford Press.Google Scholar
Rudebeck, P. H. & Murray, E. A. (2008) Amygdala and orbitofrontal cortex lesions differentially influence choices during object reversal learning. Journal of Neuroscience 28:8338–43.CrossRefGoogle ScholarPubMed
Rushworth, M. F., Buckley, M. J., Behrens, T. E., Walton, M. E. & Bannerman, D. M. (2007) Functional organization of the medial frontal cortex. Current Opinion in Neurobiology 17:220–27. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/17350820.CrossRefGoogle ScholarPubMed
Russell, J. A. (2003) Core affect and the psychological construction of emotion. Psychological Review 110:145–72.CrossRefGoogle ScholarPubMed
Russell, J. A., Bachorowski, J. & Fernandez-Dols, J. M. (2003) Facial and vocal expressions of emotion. Annual Review of Psychology 54:329–49. Available at: https://www2.bc.edu/~russeljm/publications/annurev.psych.2003.pdf.CrossRefGoogle ScholarPubMed
Russell, J. A. & Barrett, L. F. (1999) Core affect, prototypical emotional episodes, and other things called emotion: Dissecting the elephant. Journal of Personality and Social Psychology 76:805–19. Available at:http://psycnet.apa.org/journals/psp/76/5/805.html.CrossRefGoogle ScholarPubMed
Russell, J. A. & Bullock, M. (1986) On the dimensions preschoolers use to interpret facial expressions of emotion. Developmental Psychology 22:97102. Available at:http://www2.bc.edu/~russeljm/.CrossRefGoogle Scholar
Sander, D., Grafman, J. & Zalla, T. (2003) The human amygdala: An evolved system for relevance detection. Reviews in the Neurosciences 14(4):303–16. Available at:http://www.affective-sciences.org/system/files/Sander_etal_RevNeuro.PDF.CrossRefGoogle ScholarPubMed
Sander, D., Grandjean, D, Kaiser, S., Wehrle, T. & Scherer, K. R. (2007) Interaction effects of perceived gaze direction and dynamic facial expression: Evidence for appraisal theories of emotion. European Journal of Cognitive Psychology 19:470–80. Available at:http://www.lemanic-neuroscience.ch/PENSTrainingCenter/articles/dsdg_ejcp06.pdf.CrossRefGoogle Scholar
Saver, J. L. & Damasio, A. R. (1991) Preserved access and processing of social knowledge in a patient with acquired sociopathy due to ventromedial frontal damage. Neuropsychologia 29:1241–49. Available at:http://www.ncbi.nlm.nih.gov/pubmed/1791934.CrossRefGoogle Scholar
Saxe, R. & Kanwisher, N. (2003) People thinking about people: The role of the temporoparietal junction in “theory of mind.” Neuroimage 19:1835–42. Available at:http://web.mit.edu/bcs/nklab/media/pdfs/SaxeKanwisherNeuroImage03.pdf.CrossRefGoogle ScholarPubMed
Schacter, D. L., Addis, D. R. & Buckner, R. L. (2007) Remembering the past to imagine the future: The prospective brain. Nature Reviews Neuroscience 8:657–61. Available at:http://www.nature.com/nrn/journal/v8/n9/abs/nrn2213.html.CrossRefGoogle ScholarPubMed
Schachter, S. & Singer, J. (1962) Cognitive, social, and physiological determinants of an emotional state. Psychological Review 69:379–99. Available at:http://www.ncbi.nlm.nih.gov/pubmed/14497895.CrossRefGoogle ScholarPubMed
Schienle, A. & Schäfer, A. (2009) In search of specificity: Functional MRI in the study of emotional experience. International Journal of Psychophysiology 73:2226. Available at:http://www.ncbi.nlm.nih.gov/pubmed/19530276.CrossRefGoogle Scholar
Schnur, T. T., Schwartz, M. F., Kimberg, D. Y., Hirshorn, E., Coslett, H. B. & Thompson-Schill, S. L. (2009) Localizing interference during naming: Convergent neuroimaging and neuropsychological evidence for the function of Broca's area. Proceedings of the National Academy of Sciences 106:322–27. Available at:http://nihongo.j-talk.com/kanji/?link=http://www.pnas.org/content/106/1/322.full.CrossRefGoogle ScholarPubMed
Schwartz, C. E., Wright, C. I., Shin, L. M., Kagan, J., Whalen, P. J., McMullin, K. G. & Rausch, S. L. (2003) Differential amygdalar response to novel versus newly familiar neutral faces: A functional MRI probe developed for studying inhibited temperament. Biological Psychiatry 53:854–62. Available at:http://www.whalenlab.info/Publications%20Page/PDFs/Schwartz_2003.pdf CrossRefGoogle ScholarPubMed
Scott, S. K., Young, A. W., Calder, A. J., Hellawell, D. J., Aggleton, J. P. & Johnson, M. (1997) Impaired auditory recognition of fear and anger following bilateral amygdala lesions. Nature 385:254–57. Available at:http://www.cf.ac.uk/psych/home2/papers/aggleton/Nature%20-%20385%20-%20254-257.pdf CrossRefGoogle ScholarPubMed
Seeley, W. W., Menon, V., Schatzberg, A. F., Keller, J., Glover, G. H., Kenna, H., Reiss, A. L. & Greicius, M. D. (2007) Dissociable intrinsic connectivity networks for salience processing and executive control. Journal of Neuroscience 27(9):2349–56. Available at:http://neuro.cjb.net/cgi/content/abstract/27/9/2349.CrossRefGoogle ScholarPubMed
Shackman, A. J., Salomons, T. V., Slagter, H. A., Fox, A. S., Winter, J. J., Davidson, R. J. (2011) The integration of negative affect, pain, and cognitive control in the cingulate cortex. Nature Reviews Neuroscience 12:154–67. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/21331082 CrossRefGoogle ScholarPubMed
Shultz, W. (2006) Behavioral theories and the neurophysiology of reward. Annual Review of Psychology 57:87115. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/16318590.CrossRefGoogle Scholar
Siegel, A., Edinger, H. & Dotto, M. (1975) Effects of electrical stimulation of the lateral aspect of the prefrontal cortex upon attack behavior in cats. Brain Research 93:473–84.CrossRefGoogle ScholarPubMed
Siegel, A., Edinger, H. & Lowenthal, H. (1974) Effects of electrical stimulation of the medial aspect of the prefrontal cortex upon attack behavior in cats. Brain Research 66:467–79.CrossRefGoogle Scholar
Simmons, W. K., Hamann, S. B., Harenski, C. N., Hu, X. P. & Barsalou, L. W. (2008) fMRI evidence for word association and situated simulation in conceptual processing. Journal of Physiology–Paris 102:106–19. Available at:http://psychology.emory.edu/cognition/barsalou/onlinepapers.html.CrossRefGoogle ScholarPubMed
Smith, S. M., Fox, P. T., Miller, K. L., Glahn, D. C., Fox, P. M., Mackay, C. E., Filippini, N., Watkins, K. E., Toro, R., Laird, A. R. & Beckman, C. F. (2009) Correspondence of the brain's functional architecture during activation and rest. Proceedings of the National Academy of Sciences USA 106:13040–45. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed.CrossRefGoogle ScholarPubMed
Spinks, J. A., Blowers, G. H. & Shek, D. T. (1985) The role of the orienting response in the anticipation of information: A skin conductance response study. Psychophysiology 22:385–94. Available at:http://www.ncbi.nlm.nih.gov/pubmed/4023149.CrossRefGoogle ScholarPubMed
Spreng, R. N., Mar, R. A. & Kim, A. S. N. (2009) The common neural basis of autobiographical memory, prospection, navigation, theory of mind, and the default mode. A quantitative meta-analysis. Journal of Cognitive Neuroscience 21:489510.CrossRefGoogle Scholar
Sprengelmeyer, R. (2007) The neurology of disgust. Brain 130:1715–17. Available at:http://www.ncbi.nlm.nih.gov/pubmed/17550907.CrossRefGoogle ScholarPubMed
Sprengelmeyer, R., Young, A. W., Calder, A. J., Karnat, A., Lange, H., Homberg, V., Perrett, D. I. & Rowland, D. (1996) Loss of disgust. Perception of faces and emotions in Huntington's disease. Brain 119:1647–65. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed?term=Sprengelmeyer%20et%20al.%201996.CrossRefGoogle ScholarPubMed
Sprengelmeyer, R., Young, A. W., Schroeder, U., Grossenbacher, P. G., Federlein, J., Büttner, T. & Przuntek, H. (1999) Knowing no fear. Proceedings of the Royal Society of London, B: Biological Sciences 266:2451–56. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1690477/.CrossRefGoogle ScholarPubMed
Sternberg, S. (2001) Separate modifiability, mental modules, and the use of pure and composite measures to reveal them. Acta Psychologica 106:147246.CrossRefGoogle Scholar
Sutton, S. K. & Davidson, R. J. (1997) Prefrontal brain asymmetry: A biological substrate of the behavioral approach and inhibition systems. Psychological Science 8:204–10. Available at:http://www.jstor.org/stable/40063179?cookieSet=1.CrossRefGoogle Scholar
Suvak, M. K. & Barrett, L. F. (2011) Considering PTSD from the perspective of brain processes: A psychological construction analysis. Journal of Traumatic Stress 24:324. Available at:http://www.affective-science.org/pubs/2011/suvak-barrett-2011.pdf.CrossRefGoogle Scholar
Suzuki, A., Hoshino, T., Shigemasu, K. & Kawamura, M. (2006) Disgust-specific impairment of facial expression recognition in Parkinson's disease. Brain 129:707–17. Available at:http://brain.oxfordjournals.org/cgi/content/abstract/129/3/707.CrossRefGoogle ScholarPubMed
Svoboda, E., McKinnon, M. C. & Levine, B. (2006) The functional neuroanatomy of autobiographical memory: A meta-analysis. Neuropsychologia 44:2189–208. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/16806314.CrossRefGoogle ScholarPubMed
Todd, R. M. & Anderson, A. K. (2009) The neurogenetics of remembering emotions past. Proceedings of the National Academy of Sciences USA 106:18881–82.CrossRefGoogle ScholarPubMed
Tomkins, S. S. (1962) Affect imagery consciousness: Vol. 1: The positive affects. Springer.Google Scholar
Tomkins, S. S. (1963) Affect imagery consciousness: Vol. 2: The negative affects. Springer.Google Scholar
Tsakiris, M., Hesse, M. D., Boy, C., Haggard, P. & Fink, G. R. (2007) Neural signatures of body ownership: A sensory network for bodily self-consciousness. Cerebral Cortex 17:2235–44. Available at:http://cercor.oxfordjournals.org/cgi/content/full/17/10/2235.CrossRefGoogle Scholar
Tsuchiya, N., Moradi, F., Felsen, C., Yamazaki, M. & Adolphs, R. (2009) Intact rapid detection of fearful faces in the absence of the amygdala. Nature Neuroscience 12:1224–25. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2756300/.CrossRefGoogle ScholarPubMed
Urry, H. L., van Reekum, C. M., Johnstone, T., Kalin, N. H., Thurow, M. E., Schaefer, H. S., Jackson, C. A., Frye, C. J., Greischar, L. L., Alexander, A. L. & Davidson, R. J. (2006) Amygdala and ventromedial prefrontal cortex are inversely coupled during regulation of negative affect and predict the diurnal pattern of cortisol secretion among older adults. The Journal of Neuroscience 26:4415–25. Available at:http://neuro.cjb.net/cgi/content/abstract/26/16/4415.CrossRefGoogle ScholarPubMed
Van der Horst, V. G. & Holstege, G. (1998) Sensory and motor components of reproductive behavior: Pathways and plasticity. Behavior and Brain Research 92(2):157–67. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed.CrossRefGoogle ScholarPubMed
Van Snellenberg, J. X. & Wager, T. D. (2009) Cognitive and motivational functions of the human prefrontal cortex. In: Luria's Legacy in the 21st Century, ed. Goldberg, E. & Bougakov, D., pp. 3061. Oxford University Press. Available at:http://www.columbia.edu/cu/psychology/tor/Papers/luria_vans_wager_sub.pdf CrossRefGoogle Scholar
Vazdarjanova, A. & McGaugh, J. L. (1998) Basolateral amygdala is not critical for cognitive memory of contextual fear conditioning. Proceedings of the National Academy of Sciences 95:15003–5007. Available at:http://www.pnas.org/content/95/25/15003.abstract.CrossRefGoogle Scholar
Vigneau, M., Beaucousin, V., Hervé, P. Y., Duffau, H., Crivello, F., Houdé, O., Mazoyer, B. & Tzourio-Mazoyer, N. (2006) Meta-analyzing left hemisphere language areas: Phonology, semantics, and sentence processing. NeuroImage 30:1414–32. Available at:http://www.Pncbi.nlm.nih.gov/pubmed/16413796 CrossRefGoogle ScholarPubMed
Vincent, J. L., Snyder, A. Z., Fox, M. D., Shannon, B. J., Andrews, J. R., Raichle, M. E. & Buckner, R. (2006) Coherent spontaneous activity identifies a hippocampal-parietal memory network. Journal of Neurophysiology 96:3517–31. Available at:http://jn.physiology.org/cgi/content/abstract/96/6/3517.CrossRefGoogle ScholarPubMed
Vogt, B. A. (1993) Structural organization of cingulate cortex: Areas, neurons, and somatodendritic transmitter receptors. In: Neurobiology of cingulate cortex and limbic thalamus, ed. Vogt, B. A. & Gabriel, M., pp. 1970. Birkhauser.CrossRefGoogle Scholar
Vogt, B. A. (2005) Pain and emotion interactions in subregions of the cingulate gyrus. Nature Reviews Neuroscience 6:533–44.CrossRefGoogle ScholarPubMed
Vogt, B. A., Berger, G. R. & Derbyshire, S. W. (2003) Structural and functional dichotomy of human midcingulate cortex. European Journal of Neuroscience 18:3134–44. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/14656310.CrossRefGoogle ScholarPubMed
Vogt, B. A., Nimchinsky, E. A., Vogt, L. J. & Hof, P. R. (1995) Human cingulate cortex: Surface features, flat maps and cytoarchitecture. The Journal of Comparative Neurology 359:490506. Available at:http://onlinelibrary.wiley.com/doi/10.1002/cne.903590310/abstract.CrossRefGoogle ScholarPubMed
Vogt, B. A., Pandya, D. N. & Rosene, D. L. (1987) Cingulate cortex of the rhesus monkey: I. Cytoarchitecture and thalamic afferents. Journal of Comparative Neurology 26:2256–70.Google Scholar
von dem Hagen, E. A., Beaver, J. D., Ewbank, M. P., Keane, J., Passamonti, L., Lawrence, A. D. & Calder, A. J. (2009) Leaving a bad taste in your mouth but not in my insula. Social Cognitive Affective Neuroscience 4:379–86. Available at:http://www.ncbi.nlm.nih.gov/pubmed/19505971.CrossRefGoogle ScholarPubMed
Vuilleumier, P. (2005) How brains beware: Neural mechanisms of emotional attention. Trends in Cognitive Sciences 12:585–94. Available at:http://www.ncbi.nlm.nih.gov.ezp-prod1.hul.harvard.edu/pubmed/16289871.CrossRefGoogle Scholar
Vuilleumier, P. & Driver, J. (2007) Modulation of visual processing by attention and emotion: Windows on causal interactions between human brain regions. Philosophical Transactions of the Royal Society B: Biological Sciences 362:837–55. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2430001/.CrossRefGoogle ScholarPubMed
Vytal, K. & Hamann, S. (2010) Neuroimaging support for discrete neural correlates of basic emotions: A voxel-based meta-analysis. Journal of Cognitive Neuroscience 22(12):2864–85.CrossRefGoogle ScholarPubMed
Wager, T., Barrett, L. F., Bliss-Moreau, E., Lindquist, K. A., Duncan, S., Kober, H., Joseph, J., Davidson, M. & Mize, J. (2008) The neuroimaging of emotion. In: Handbook of emotions, 3rd edition. Guilford Press.Google Scholar
Wager, T. D., Jonides, J., Smith, E. E. & Nichols, T. E. (2005) Towards a taxonomy of attention-shifting: Individual differences in fMRI during multiple shift types. Cognitive Affective and Behavioral Neuroscience 5:127–43. Available at:http://psych.colorado.edu/~tor/Papers/Wager_2005_CABN_shifting_indivdiffs_fmri.pdf.CrossRefGoogle Scholar
Wager, T. D., Lindquist, M. & Kaplan, L. (2007) Meta-analysis of functional neuroimaging data: Current and future directions. Social Cognitive and Affective Neuroscience 2:150–58. Available at:http://scan.oxfordjournals.org/content/2/2/150.abstract.CrossRefGoogle ScholarPubMed
Wager, T. D., Phan, K. L., Liberzon, I. & Taylor, S. F. (2003) Valence, gender, and lateralization of functional brain anatomy in emotion: A meta-analysis of findings from neuroimaging. NeuroImage 19:513–31. Available at:http://www.ncbi.nlm.nih.gov/pubmed/12880784.CrossRefGoogle ScholarPubMed
Wager, T. D., Reading, S. & Jonides, J. (2004) Neuroimaging studies of shifting attention: A meta-analysis. NeuroImage 22:1679–93. http://psych.colorado.edu/~tor/Papers/Wager_2004_Switching_Meta.pdf.CrossRefGoogle ScholarPubMed
Wager, T. D. & Smith, E. E. (2003) Neuroimaging studies of working memory: A meta-analysis. Cognitive, Affective and Behavioral Neuroscience 3:255–74. Available at:http://www.columbiauniversity.org/cu/psychology/tor/Papers/Wager_Smith_2003_WM_meta.pdf.CrossRefGoogle ScholarPubMed
Wager, T. D., van Ast, V. A., Hughes, B. L., Davidson, M. L., Lindquist, M. A. & Ochsner, K. N. (2009a) Brain mediators of cardiovascular responses to social threat, Part II: Prefrontal-subcortical pathways and relationship with anxiety. NeuroImage 47:836–51.CrossRefGoogle ScholarPubMed
Wager, T. D., Waugh, C. E., Lindquist, M., Noll, D. C., Fredrickson, B. L. & Taylor, S. F. (2009b) Brain mediators of cardiovascular responses to social threat, Part I: Reciprocal dorsal and ventral sub-regions of the medial prefrontal cortex and heart-rate reactivity. Neuroimage 47:821–35.CrossRefGoogle ScholarPubMed
Wang, G., Tomasi, D., Backus, W., Wang, R., Telang, F., Geliebter, A., Korner, J., Bauman, A., Fowler, J. S., Thanos, P. K. & Volknow, N. D. (2008) Gastric distention activates satiety circuitry in the human brain. NeuroImage 39:1824–31. Available at:http://www.ncbi.nlm.nih.gov/pubmed/18155924.CrossRefGoogle ScholarPubMed
Warnick, J. E., LaPorte, J. L. & Kalueff, A. V. (2010) Domain interplay in mice and men: New possibilities for the “Natural Kinds” theory of emotion. New Ideas in Psychology 29:4956.CrossRefGoogle Scholar
Weierich, M. R., Wright, C. I., Negreira, A., Dickerson, B. C. & Barrett, L. F. (2010) Novelty as a dimension in the affective brain. NeuroImage 49:2871–78. http://www.ncbi.nlm.nih.gov/pubmed/19796697.CrossRefGoogle ScholarPubMed
Whalen, P. J. (2007) The uncertainty of it all. Trends in Cognitive Sciences 11:499500. Available at:http://www.whalenlab.info/Publications%20Page/PDFs/Whalen_TICS_2007.pdf.CrossRefGoogle Scholar
Whalen, PJ. (1998) Fear, vigilance, and ambiguity: Initial neuroimaging studies of the human amygdala. Current Directions in Psychological Science 7:177188. Available at:http://whalenlab.info/publications.html CrossRefGoogle Scholar
Whalen, P. J., Kagan, J., Cook, R. G., Davis, F. C., Kim, H., Polis, S., McLaren, D., Somerville, L. H., McLean, A. A., Maxwell, J. S. & Johnstone, T. (2004) Human amygdala responsivity to masked fearful eye whites. Science 306:2061. Available at:http://www.whalenlab.info/Publications%20Page/PDFs/Whalen_etal_Sc_2004.pdf.CrossRefGoogle ScholarPubMed
Whalen, P. J., Rauch, S. L., Etcoff, N. L., McInerney, S. C., Lee, M. B. & Jenike, M. A. (1998) Masked presentations of emotional facial expressions modulate amygdala activity without explicit knowledge. Journal of Neuroscience 18:411. Available at:http://neuro.cjb.net/cgi/content/abstract/18/1/411.CrossRefGoogle ScholarPubMed
Whalen, P. J., Shin, L. M., McInerney, S. C., Fischer, H., Wright, C. I. & Rauch, S. L. (2001) A functional MRI study of human amygdala responses to facial expressions of fear vs. anger. Emotion 1:7083. Available at:http://www.whalenlab.info/Publications%20Page/PDFs/Whalen_Shin_2001.pdf.CrossRefGoogle Scholar
Wicker, B., Keysers, C., Plailly, J., Royet, J. P., Gallese, V. & Rizzolati, G. (2003) Both of us disgusted in my insula: The common neural basis of seeing and feeling disgust. Neuron 40:655–64. Available at:http://www.ncbi.nlm.nih.gov/pubmed/14642287.CrossRefGoogle Scholar
Wilson, F. A. W. & Rolls, E. T. (1993) The effects of stimulus novelty and familiarity on neuronal activity in the amygdala of monkeys performing recognition memory tasks. Experimental Brain Research 93:367–82. Available at:http://www.springerlink.com/content/g222263510052042/.CrossRefGoogle ScholarPubMed
Wilson-Mendenhall, C. D., Barrett, L. F., Simmons, W. K. & Barsalou, L. W. (2011) Grounding emotion in situated conceptualization. Neuropsychologia 49:1105–27. Available at:http://psychology.emory.edu/cognition/barsalou/papers/Wilson_Mendenhall_et_al-Neuropsychologia_in_press-grounding_emotion.pdf.CrossRefGoogle ScholarPubMed
Wright, C. I., Fischer, H., Whalen, P. J., McInerney, S. C., Shin, L. M. & Rauch, S. L. (2001) Differential prefrontal cortex and amygdala habituation to repeatedly presented emotional stimuli. NeuroReport 12:379–83. Available at:http://journals.lww.com/neuroreport/Abstract/2001/02120/Differential_prefrontal_cortex_and_amygdala.39.aspx.CrossRefGoogle ScholarPubMed
Wright, C. I., Martis, B., Schwartz, C. E., Shin, L. M., Fischer, H. H., McMullin, K. & Rausch, S. (2003) Novelty responses and differential effects of order in the amygdala, substantia innominata, and inferior temporal cortex. NeuroImage 18:660–69. Available at:http://www.ncbi.nlm.nih.gov/pubmed/12667843.CrossRefGoogle ScholarPubMed
Wright, C. I., Negreira, A., Gold, A. L., Britton, J. C., Williams, D. & Barrett, L. F. (2008) Neural correlates of novelty and face-age effects in young and elderly adults. NeuroImage 42:956–68. Available at:http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2613685/.CrossRefGoogle Scholar
Wright, C. I., Wedig, M. M., Williams, D., Rauch, S. L. & Albert, M. S. (2006) Novel fearful faces activate the amygdala in healthy young and elderly adults. Neurobiology of Aging 27:361–74. Available at:http://www.neurobiologyofaging.org/article/S0197-4580%2805%2900069-2/abstract.CrossRefGoogle Scholar
Wundt, W. (1897/1998) Outlines of psychology, trans. Judd, C. H.. Thoemmes Press. (Original work published in 1897).CrossRefGoogle Scholar
Xu, F. (2002) The role of language in acquiring object kind concepts in infancy. Cognition 85:223–50. Available at:http://babylab.berkeley.edu/ProfFeiXu.html.CrossRefGoogle ScholarPubMed
Yang, Y. & Raine, A. (2009) Neuroimaging: Prefrontal structural and functional brain imaging findings in antisocial, violent, and psychopathic individuals: A meta-analysis. Psychiatry Research 174:8188. Available at:http://www.ncbi.nlm.nih.gov/pubmed/19833485.CrossRefGoogle ScholarPubMed
Zahn, R., Moll, J., Paiva, M., Garrido, G., Krueger, F., Huey, E. D. & Grafman, J. (2009) The neural basis of human social values: Evidence from functional MRI. Cerebral Cortex 19:276–83. Available at: http://cercor.oxfordjournals.org/cgi/content/abstract/19/2/276.CrossRefGoogle ScholarPubMed
Figure 0

Figure 1. Locationist Hypotheses of Brain–Emotion Correspondence. A: Lateral view. B: Sagital view at the midline. C: Ventral view. D: Coronal view. Brain regions hypothesized to be associated with emotion categories are depicted. Here we depict the most popular locationist hypotheses, although other locationist hypotheses of brain–emotion correspondence exist (e.g., Panksepp, 1998). Fear: amygdala (yellow); Disgust: insula (green); Anger: OFC (rust); Sadness: ACC (blue). A color version of this image can be viewed in the online version of this target article at http://www.journals.cambridge.org/bbs.

Figure 1

Figure 2. Psychological Constructionist Hypotheses of Brain–Emotion Correspondence. A: Lateral view. B: Sagital view at the midline. C: Ventral view. D: Coronal view. Brain regions hypothesized to be associated with psychological operations are depicted. In some cases, we present only the key brain regions within networks that have been empirically linked to our hypothesized psychological operations. In instances where the whole brain network is not depicted, we point readers to relevant literature. Core Affect (pink): amygdala, insula, mOFC (Bas 10m, 11m, 13a, b, 14r, c), lOFC (BAs 47, 12, 13l, m, 11l), ACC (Bas, 32, 24, 25), thalamus, hypothalamus, bed nucleus of the stria terminalis, basal forebrain, PAG. Conceptualization (purple): VMPFC (Bas 11, 25, 32, 34), DMPFC (BAs 9, 10p), medial temporal lobe* (hippocampus, entorhinal cortex, parahippocampal cortex), posterior cingulate cortex/retrosplenial area (BA 23, 31). Language (green): VLPFC (Bas 44, 45, 46), anterior temporal lobe (BA 38); for additional regions, see Vigneau et al. (2006). Executive Attention (orange): DLPFC (BAs 9, 10, 46), VLPFC (BAs 44, 45, 46); for additional regions see Corbetta & Shulman, (2002); Dosenbach et al. (2006); Wager et al. (2004). (*this structure is not visible in this view of the brain). A color version of this image can be viewed in the online version of this target article at http://www.journals.cambridge.org/bbs.

Figure 2

Figure 3. Kober et al.'s (2008) Functional Clusters. Kober et al.'s (2008) six functional clusters are consistent with the ingredients hypothesized by our psychological constructionist model. The brain areas making up the “core limbic group” and “lateral paralimbic group” are part of the network that helps to constitute core affect. Aspects of the “medial posterior group” and “medial PFC group” are part of the network involved with conceptualization. Areas in the “cognitive/motor control group” are consistent with the networks supporting language and executive attention. In addition, an “occipital/visual group” was also identified as part of the neural reference space for emotion. Visual cortex has connectivity with areas involved in core affect (e.g., amygdala, orbitofrontal cortex; Amaral & Price 1984; Barrett & Bar 2009; Pessoa & Adolphs 2010), and there is growing evidence that a person's core affective state modulates activity in primary visual cortex (Damaraju et al. 2009). Core affect even shapes aspects of visual perception ranging from contrast sensitivity (Phelps et al. 2006) to visual awareness (Anderson et al. 2011). A color version of this image can be viewed in the online version of this target article at http://www.journals.cambridge.org/bbs.

Figure 3

Figure 4. The Neural Reference Space for Discrete Emotion. The neural reference space (phrase coined by Edelman [1989]) is the set of brain regions consistently activated across all studies assessing the experience or perception of anger, disgust, fear, happiness and sadness (i.e. the superordinate category emotion). Brain regions in yellow exceeded the height threshold (p<05) and regions in orange exceeded the most stringent extent-based threshold (p<001). Regions in pink and magenta correspond to lesser extent-based thresholds and are not discussed in this article. Cortex is grey, the brainstem and nucleus accumbens are green, the amygdala is blue and the cerebellum is purple. A color version of this image can be viewed in the online version of this target article at http://www.journals.cambridge.org/bbs.

Figure 4

Figure 5. Logistic Regression Findings. Selected results from the logistic regressions are presented (for additional findings, see Table S6 in supplementary materials). Circles with positive values represent a 100% increase in the odds that a variable predicted an increase in activity in that brain area. Circles with negative values represent a 100% increase in the odds that a variable predicted there would not be an increase in activity in that brain area. Legend: Blue lines: left hemisphere; Green lines: right hemisphere. Arrowheads: % change in odds is greater than values represented in this figure. Abbreviations: OFC: orbitofrontal cortex; DLPFC: dorsolateral prefrontal cortex; ATL: anterior temporal lobe; VLPFC: ventrolateral prefrontal cortex; DMPFC: dorsomedial prefrontal cortex; aMCC: anterior mid-cingulate cortex; sAAC: subgenual ACC. A color version of this image can be viewed in the online version of this target article at http://www.journals.cambridge.org/bbs.

Figure 5

Figure 6. Proportion of Study Contrasts with Increased Activation in Four Key Brain Areas. The y-axes plot the proportion of study contrasts in our database that had increased activation within 10mm of that brain area. The x-axes denote the contrast type separated by experience (exp) and perception (per). All brain regions depicted are in the right hemisphere. See Figures S2 and S3 in supplementary materials, available at http://www.journals.cambridge.org/bbs2012008, for additional regions. A color version of this image can be viewed in the online version of this target article at http://www.journals.cambridge.org/bbs.

Figure 6

Table 1. Brain Regions with a Consistent Increase in Activity Associated with the Experience or Perception of Discrete Emotion Categories in Density Analyses

Figure 7

Table 2. Brain Regions with a Consistent Increase in Activity Associated with the Experience or Perception of Discrete Emotion Categories in χ2 Analyses

Figure 8

Table 3. Summary of Brain Regions Showing Consistent Increases in Activation During Mental States and Methodological Manipulations

Supplementary material: File

Lindquist et al. supplementary material

Supplementary table and figures

Download Lindquist et al. supplementary material(File)
File 1.2 MB
Supplementary material: File

Lindquist et al. supplementary material

Supplementary references

Download Lindquist et al. supplementary material(File)
File 61.4 KB