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Journal Article•DOI•

Brain mechanisms for emotional influences on perception and attention: What is magic and what is not

Gilles Pourtois1, Antonio Schettino1, Patrik Vuilleumier2•
Ghent University1, University of Geneva2
01 Mar 2013-Biological Psychology (Elsevier Publishing)-Vol. 92, Iss: 3, pp 492-512
TL;DR: It is suggested that emotion signals may enhance processing efficiency and competitive strength of emotionally significant events through gain control mechanisms similar to those of other attentional systems, but mediated by distinct neural mechanisms in amygdala and interconnected prefrontal areas.
About: This article is published in Biological Psychology.The article was published on 2013-03-01 and is currently open access. It has received 577 citations till now. The article focuses on the topics: Event-related potential & Perception.

Summary (2 min read)

Jump to: [Introduction] – [Neural routes to the amygdala] and [Conclusions and perspectives]

Introduction

  • 2 The rapid and efficient selection of emotionally-salient or goal-relevant stimuli in the environment is crucial for flexible and adaptive behaviors.
  • Converging data from neuroscience and psychology have accrued during the last decade to identify brain systems involved in emotion processing, selective attention, and their interaction, which together act to extract the emotional or motivational value of sensory events and respond appropriately.
  • An important hub in these systems is the amygdala, which may not only monitor the emotional value of stimuli, but also readily project to several other areas and send feedback to sensory pathways (including striate and extrastriate visual cortex).
  • This system generates saliency signals that modulate perceptual, motor, as well as memory processes, and thus in turn regulate behavior appropriately.
  • Here, the authors review their current views on the function and properties of these brain systems, with an emphasis on their involvement in the rapid and/or preferential processing of threat-relevant stimuli.

Neural routes to the amygdala

  • The fact that some emotional processing may take place in the amygdala without voluntary attention (or sometimes without awareness) has raised questions about the possible pathways conveying information to this region, prior to full cortical processing.
  • These results support a dual route hypothesis, but a subcortical route or cortico-cortical long-range pathways appear equally plausible, and the exact anatomical substrates of these effects therefore remain to be clarified.
  • A role for the amygdala and emotional influences on attention in these tasks is supported by the convergence of these behavioral effects in healthy participants with patterns of neurophysiological responses in imaging studies, as well as observations in patients with lesions to the amygdala.
  • These data therefore suggest that amygdala damage may have an important impact on attention to salient face information, rather than on the recognition of fear cues per se.

Conclusions and perspectives

  • These modulatory effects of emotion closely resemble the typical enhancement produced in the same sensory regions by selective attention mechanisms, and normally associated with top-down signals from dorsal fronto-parietal networks (Desimone & Duncan, 1995; Kastner & Ungerleider, 2000; Posner & Dehaene, 1994; Corbetta & Shulman, 2002).
  • In sum, therefore, there is no reason to assume that reflexive (i.e. automatic and involuntary) emotional responses in amygdala and subsequent influences on perception would imply a process that is totally encapsulated, “resource” free, and unaffected by changes in amount or duration of sensory inputs (e.g. Pessoa, Kastner, et al., 2002; Pessoa, 2005).
  • This focus of past research is understandable given the obvious and vital importance of efficient attention to threat (see Compton, 2003; Vuilleumier, 2005), as well as the crucial links with amygdala function (LeDoux, 1996; Amaral, et al., 2003; Phelps & LeDoux, 2005) and the direct implications for psychopathology (Öhman & Mineka, 2001; Bar-Haim, et al., 2007).
  • By doing so, researchers should go beyond the traditional notions of a single attention system or binary (e.g. controlled vs. automatic) divisions, but instead redefine the corresponding psychological construct and behavioral phenomena in terms of their underlying neural processes.

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Citations
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Journal Article•DOI•
Coming to terms with fear

[...]

Joseph E. LeDoux1•
Center for Neural Science1
25 Feb 2014-Proceedings of the National Academy of Sciences of the United States of America
TL;DR: A conception of so-called fear conditioning in terms of circuits that operate nonconsciously, but that indirectly contribute to conscious fear, is proposed as way forward.
Abstract: The brain mechanisms of fear have been studied extensively using Pavlovian fear conditioning, a procedure that allows exploration of how the brain learns about and later detects and responds to threats. However, mechanisms that detect and respond to threats are not the same as those that give rise to conscious fear. This is an important distinction because symptoms based on conscious and nonconscious processes may be vulnerable to different predisposing factors and may also be treatable with different approaches in people who suffer from uncontrolled fear or anxiety. A conception of so-called fear conditioning in terms of circuits that operate nonconsciously, but that indirectly contribute to conscious fear, is proposed as way forward.

678 citations

Journal Article•DOI•
Using Neuroscience to Help Understand Fear and Anxiety: A Two-System Framework.

[...]

Joseph E. LeDoux1, Daniel S. Pine2•
Center for Neural Science1, Nathan Kline Institute for Psychiatric Research2
09 Sep 2016-American Journal of Psychiatry
TL;DR: It is argued that failure to recognize and consistently emphasize a distinction between circuits underlying two classes of responses elicited by threats has impeded progress in understanding fear and anxiety disorders and hindered attempts to develop more effective pharmaceutical and psychological treatments.
Abstract: Tremendous progress has been made in basic neuroscience in recent decades. One area that has been especially successful is research on how the brain detects and responds to threats. Such studies have demonstrated comparable patterns of brain-behavior relationships underlying threat processing across a range of mammalian species, including humans. This would seem to be an ideal body of information for advancing our understanding of disorders in which altered threat processing is a key factor, namely, fear and anxiety disorders. But research on threat processing has not led to significant improvements in clinical practice. The authors propose that in order to take advantage of this progress for clinical gain, a conceptual reframing is needed. Key to this conceptual change is recognition of a distinction between circuits underlying two classes of responses elicited by threats: 1) behavioral responses and accompanying physiological changes in the brain and body and 2) conscious feeling states reflected in sel...

598 citations

Journal Article•DOI•
Norepinephrine ignites local hotspots of neuronal excitation: How arousal amplifies selectivity in perception and memory.

[...]

Mara Mather1, David Clewett1, Michiko Sakaki2, Carolyn W. Harley3•
University of Southern California1, University of Reading2, Memorial University of Newfoundland3
01 Jan 2016-Behavioral and Brain Sciences
TL;DR: GANE not only reconciles apparently contradictory findings in the emotion-cognition literature but also extends previous influential theories of LC neuromodulation by proposing specific mechanisms for how LC-NE activity increases neural gain.
Abstract: Emotional arousal enhances perception and memory of high-priority information but impairs processing of other information. Here, we propose that, under arousal, local glutamate levels signal the current strength of a representation and interact with norepinephrine (NE) to enhance high priority representations and out-compete or suppress lower priority representations. In our "glutamate amplifies noradrenergic effects" (GANE) model, high glutamate at the site of prioritized representations increases local NE release from the locus coeruleus (LC) to generate "NE hotspots." At these NE hotspots, local glutamate and NE release are mutually enhancing and amplify activation of prioritized representations. In contrast, arousal-induced LC activity inhibits less active representations via two mechanisms: 1) Where there are hotspots, lateral inhibition is amplified; 2) Where no hotspots emerge, NE levels are only high enough to activate low-threshold inhibitory adrenoreceptors. Thus, LC activation promotes a few hotspots of excitation in the context of widespread suppression, enhancing high priority representations while suppressing the rest. Hotspots also help synchronize oscillations across neural ensembles transmitting high-priority information. Furthermore, brain structures that detect stimulus priority interact with phasic NE release to preferentially route such information through large-scale functional brain networks. A surge of NE before, during, or after encoding enhances synaptic plasticity at NE hotspots, triggering local protein synthesis processes that enhance selective memory consolidation. Together, these noradrenergic mechanisms promote selective attention and memory under arousal. GANE not only reconciles apparently contradictory findings in the emotion-cognition literature but also extends previous influential theories of LC neuromodulation by proposing specific mechanisms for how LC-NE activity increases neural gain.

458 citations

Cites background from "Brain mechanisms for emotional infl..."

  • ...For example, in their multiple attention gain control (MAGiC) model, Pourtois et al. (2013) argue that emotional salience shapes perception via amplification mechanisms independent of other attentional processes....

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Journal Article•DOI•
Exogenous (automatic) attention to emotional stimuli: a review

[...]

Luis Carretié1•
Autonomous University of Madrid1
29 Mar 2014-Cognitive, Affective, & Behavioral Neuroscience
TL;DR: The present review reveals that there are several key issues for which experimental data are surprisingly scarce, and confirms the relevance of including emotional distractors in studies on exogenous attention.
Abstract: Current knowledge on the architecture of exogenous attention (also called automatic, bottom-up, or stimulus-driven attention, among other terms) has been mainly obtained from studies employing neutral, anodyne stimuli. Since, from an evolutionary perspective, exogenous attention can be understood as an adaptive tool for rapidly detecting salient events, reorienting processing resources to them, and enhancing processing mechanisms, emotional events (which are, by definition, salient for the individual) would seem crucial to a comprehensive understanding of this process. This review, focusing on the visual modality, describes 55 experiments in which both emotional and neutral irrelevant distractors are presented at the same time as ongoing task targets. Qualitative and, when possible, meta-analytic descriptions of results are provided. The most conspicuous result is that, as confirmed by behavioral and/or neural indices, emotional distractors capture exogenous attention to a significantly greater extent than do neutral distractors. The modulatory effects of the nature of distractors capturing attention, of the ongoing task characteristics, and of individual differences, previously proposed as mediating factors, are also described. Additionally, studies reviewed here provide temporal and spatial information—partially absent in traditional cognitive models—on the neural basis of preattention/evaluation, reorienting, and sensory amplification, the main subprocesses involved in exogenous attention. A model integrating these different levels of information is proposed. The present review, which reveals that there are several key issues for which experimental data are surprisingly scarce, confirms the relevance of including emotional distractors in studies on exogenous attention.

304 citations

Cites background from "Brain mechanisms for emotional infl..."

  • ...…to emotional stimuli and in modulating attentional resources to them is consistently supported (see reviews in Adolphs & Spezio, 2007; Phelps, 2005; Pourtois et al., 2012; Vuilleumier, 2005; Wager, Phan, Liberzon & Taylor, 2003), its mandatory or central involvement (or at least, its higher…...

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  • ...Early emotional effects (C1 component, peaking around 90– 100 ms) have also been observed in tasks other than CDTD (see a review in Pourtois et al., 2012)....

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  • ...Enhanced responses in the visual cortex to emotional visual stimuli, as compared with neutral stimuli, have been often and consistently reported (see reviews in Carretié, Albert, et al., 2009; Pourtois et al., 2012)....

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  • ...While the crucial role of the human amygdala in organizing the response to emotional stimuli and in modulating attentional resources to them is consistently supported (see reviews in Adolphs & Spezio, 2007; Phelps, 2005; Pourtois et al., 2012; Vuilleumier, 2005; Wager, Phan, Liberzon & Taylor, 2003), its mandatory or central involvement (or at least, its higher hierarchy) in preattentional evaluation of the environment is currently under debate....

    [...]

Journal Article•DOI•
The neurobiology of emotion-cognition interactions: fundamental questions and strategies for future research.

[...]

Hadas Okon-Singer1, Talma Hendler2, Luiz Pessoa3, Alexander J. Shackman3•
University of Haifa1, Tel Aviv University2, University of Maryland, College Park3
17 Feb 2015-Frontiers in Human Neuroscience
TL;DR: This research demonstrates that stress, anxiety, and other kinds of emotion can profoundly influence key elements of cognition, including selective attention, working memory, and cognitive control, and suggests that widely held beliefs about the key constituents of ‘the emotional brain’ and “the cognitive brain” are fundamentally flawed.
Abstract: Recent years have witnessed the emergence of powerful new tools for assaying the brain and a remarkable acceleration of research focused on the interplay of emotion and cognition. This work has begun to yield new insights into fundamental questions about the nature of the mind and important clues about the origins of mental illness. In particular, this research demonstrates that stress, anxiety, and other kinds of emotion can profoundly influence key elements of cognition, including selective attention, working memory, and cognitive control. Often, this influence persists beyond the duration of transient emotional challenges, partially reflecting the slower molecular dynamics of catecholamine and hormonal neurochemistry. In turn, circuits involved in attention, executive control, and working memory contribute to the regulation of emotion. The distinction between the ‘emotional’ and the ‘cognitive’ brain is fuzzy and context-dependent. Indeed, there is compelling evidence that brain territories and psychological processes commonly associated with cognition, such as the dorsolateral prefrontal cortex and working memory, play a central role in emotion. Furthermore, putatively emotional and cognitive regions influence one another via a complex web of connections in ways that jointly contribute to adaptive and maladaptive behavior. This work demonstrates that emotion and cognition are deeply interwoven in the fabric of the brain, suggesting that widely held beliefs about the key constituents of ‘the emotional brain’ and ‘the cognitive brain’ are fundamentally flawed. We conclude by outlining several strategies for enhancing future research. Developing a deeper understanding of the emotional-cognitive brain is important, not just for understanding the mind but also for elucidating the root causes of its disorders.

279 citations

Cites background from "Brain mechanisms for emotional infl..."

  • ...…in Human Neuroscience www.frontiersin.org February 2015 | Volume 9 | Article 58 | 1 (e.g., Siman-Tov et al., 2009; Lerner et al., 2012; Pourtois et al., 2013; Carretié, 2014) the ability to selectively respond to relevant aspects of the environment while inhibiting potential sources…...

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  • ...Altogether, these results converge to support a multiple systems architecture responsible for attention control (Corbetta and Shulman, 2002; Vuilleumier, 2005), wherein the effects of emotional attention may flexibly add to those imposed by other bottom–up or top–down control processes....

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Abstract: The two basic phenomena that define the problem of visual attention can be illustrated in a simple example. Consider the arrays shown in each panel of Figure 1. In a typical experiment, before the arrays were presented, subjects would be asked to report letters appearing in one color (targets, here black letters), and to disregard letters in the other color (nontargets, here white letters). The array would then be briefly flashed, and the subjects, without any opportunity for eye movements, would give their report. The display mimics our. usual cluttered visual environment: It contains one or more objects that are relevant to current behavior, along with others that are irrelevant. The first basic phenomenon is limited capacity for processing information. At any given time, only a small amount of the information available on the retina can be processed and used in the control of behavior. Subjectively, giving attention to any one target leaves less available for others. In Figure 1, the probability of reporting the target letter N is much lower with two accompa­ nying targets (Figure la) than with none (Figure Ib). The second basic phenomenon is selectivity-the ability to filter out un­ wanted information. Subjectively, one is aware of attended stimuli and largely unaware of unattended ones. Correspondingly, accuracy in identifying an attended stimulus may be independent of the number of nontargets in a display (Figure la vs Ie) (see Bundesen 1990, Duncan 1980).

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Harvard University1, Yale University2
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"Brain mechanisms for emotional infl..." refers background or methods in this paper

  • ...This does not preclude that different task demands or voluntary emotion regulation strategies can alter (increase or decrease) this readiness to respond (Ochsner and Gross, 2005; Vrticka et al., 2011), possibly through direct modulation of amygdala circuitry and/or indirect modulation of its inputs....

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  • ...4; Drevets and Raichle, 998; Ochsner and Gross, 2005; Etkin et al., 2010a), but also hrough changes in the “firing threshold” settings of amygdala ciruitry itself....

    [...]

  • ...…emotion regulation strategies, as well as personality factors, genetic makeup, and specific psychiatric disorders such as anxiety and depression (e.g. Drevets, 2003; Ochsner and Gross, 2005; Etkin et al., 2010a; Cornwell et al., 2011; Pezawas et al., 2005; Hariri et al., 2002; Canli et al., 2002)....

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  • ...For example, the motional attention system might be either amplified or attenated by top–down modulations from higher-order regions in edial prefrontal/anterior cingulate/orbitofrontal regions involved n emotion regulation processes (see Fig. 4; Drevets and Raichle, 998; Ochsner and Gross, 2005; Etkin et al., 2010a), but also hrough changes in the “firing threshold” settings of amygdala ciruitry itself....

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  • ...…network controlling amygdala responsiveness, then the use of explicit regulation strategies such as self-distancing or reappraisal (see Ochsner and Gross, 2005; Kalisch, 2009) may turn out to be difficult to implement, and therefore less effective than manipulations acting on “intact”…...

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Frequently Asked Questions (9)
Q1. What is the effect of emotional faces on the visual field?

In addition, due to the complexity of emotional cues, some effects obtained with emotional scenes tend to occur later than those evoked by faces and show longer duration (and more sustained effects), such as the EPN and LPP (Schupp, et al., 2006; Sabatinelli, et al., 2007; Hajcak, et al., in press). 

the loss of emotional effect predominated in visual areas on the same side as the amygdala lesions, suggesting a causal impact on distant visual areas caused by intrahemispheric influences, in keeping with the direct feedback connections from amygdala established by anatomical studies (Amaral, et al., 2003). 

While the amygdala might be critical to learn the emotional significance of novel stimuli and respond to simple visual cues associated with emotional relevance (as can be conveyed by quick and coarse sensory pathways), differential reactions to some over-learned as well as more complex stimuli might require additional processes, possibly mediated by higher sensory (e.g. extrastriate) cortices or posterior orbitofrontal regions receiving inputs from both sensory regions and amygdala (Barbas, Zikopoulos, & Timbie, 2010).33Data from neuroscience reviewed in this paper point to the existence of brain mechanisms centered on the amygdala and interconnected areas (OFC, cholinergic nuclei), whose main function is to assess the emotional value of sensory events and boost their perceptual processing in early sensory pathways, presumably allowing the organism to swiftly respond to emotionallyrelevant stimuli in the environment and extract relevant information about their nature and location (Vuilleumier, 2005; Vuilleumier & Huang, 2009). 

These findings suggest a source of modulation by emotion cues that is independent from voluntary attention and/or produces involuntary shifts in selective attention. 

Projections to early visual areas may also account for increased activations in V1/V2 (Lang, et al., 1998; Pessoa, McKenna, et al., 2002) and extrastriate areas in occipital cortex (Lane, et al., 1997; Sabatinelli, et al., 2005; Sabatinelli, et al., 2007), with corresponding effects of the C1 and P1 components in early visual ERP. 

the enhancement of P1 evoked by an emotional face has also been found to predict the magnitude of covert expression mimicry produced by the viewer (as indexed by facial EMG; Achaibou, Pourtois, Schwartz, & Vuilleumier, 2008), a result indicating that this early perceptual enhancement might also contribute to the recognition of emotional expressions and influence motor behavior. 

The brief presentation of a (task-irrelevant) upright fearful face (as opposed to either a neutral face or an inverted fearful face) enhances visual sensitivity for the orientation of a subsequently presented low-spatial frequency stimulus (such as a Gabor patch; Phelps, et al., 2006; Bocanegra & Zeelenberg, 2009). 

Each of the three cueing effects was found to contribute to spatial orienting of attention and combined in an additive manner to facilitate target detection and reaction times (see Fig. 1B). 

This phenomenon has been extensively demonstrated by neuronal recordings as well as imaging methods (EEG, PET, fMRI), and attributed to a gain control mechanism exerted by a fronto-parietal network (see Posner & Dehaene, 1994; Corbetta & Shulman, 2002) on lower-level sensory areas that can enhance the relevant/attended stimulus representation, while suppressing the irrelevant/unattended stimulus representation (see Fig.