(Oxford Library of Psychology) Theodore P. Beauchaine, Sheila E. Crowell - The Oxford Handbook of Emotion Dysregulation-Oxford University Press (2020)
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Te Oxford Handbook of Emotion Dysregulation
Oxford Library of Psy Psychology chology Area Editors: Clinical Psychology David H. Barlow Cognitive Neuroscience Kevin N. Ochsner and Stephen M. Kosslyn Cognitive Psychology Daniel Reisberg Counseling Psychology Elizabeth M. Altmaier and Jo-Ida C. Hansen Developmental Psychology Philip David Zelazo Health Psychology Howard S. Friedman History of Psychology David B. Baker Methods and Measurement Measurement Todd D. Little Neuropsychology Kenneth M. Adams Organizational Psychology Steve W. J. Kozlowski Personality and Social Psychology Personality Kay Deaux and Mark Snyder
O X F O R D
L I B R A R Y
O F
P S Y C H O L O G Y
Te Oxford Handbook of Emotion Dysregulation Edited by
Teodore P. Beauchaine Sheila E. Crowell
1
1 Oxford University Press is a department of the University of Oxford. It furthers the University’s University’s objective of excellence in research, scholarship, and education e ducation by publishing worldwide. Oxford is a registered trade mark of Oxford University Press in the UK and certain other countries. Published in the United States of America by Oxford University Press Madison Avenue, New York, York, NY , United States of America. © Oxford University Press 2020 All rights reserved. Notransmitted, part of thisinpublication may storedthe in a retrieval system, or any form or bybe anyreproduced, means, without prior permission in writing of Oxford University Press, or as expressly permitted by law, by license, or under terms agreed with the appropriate reproduction rights organization. Inquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above. You Y ou must not circulate this work in any other form and you must impose this same condition on any acquirer. Library of Congress Cataloging-in-Publication Data Names: Beauchaine, Teodore P., editor. | Crowell, Sheila E. (Sheila Elizabeth), editor. itle: Te Oxford handbook of emotion dysregulation / edited by Teodore P. P. Beauchaine, Sheila E. Crowell. Description: New York, York, NY : Oxford University Press, [2020] | Series: Oxford library of psychology | Includes bibliographical references and index. | Identifiers: LCCN 2019031334 (print) | LCCN 2019031335 (ebook) | ISBN 9780190689285ISBN (hardback) | ISBN 9780190689308 (epub) | 9780190689292 Subjects: LCSH: Emotions—Handbooks, manuals, etc. | Psychology, Pathological— Handbooks, manuals, etc. | Affect (Psychology)—Handbooks, manuals, etc. Classification: LCC RC455.4.E46 RC455.4.E46 O94 2020 (print) | LCC RC455.4.E46 (ebook) | DDC 616.89—dc23 LC record available at https://lccn.loc.gov/2019031334 LC ebook record available at https://lccn.loc.gov/2019031335 9 8 7 6 5 4 3 2 1 Printed by Integrated Books International, United States of America
CONTRIBUTORS
Molly Adrian, PhD
Patricia A. Brennan, PhD
Department of Psychiatry and Behavioral Sciences Seattle Children’s Hospital Kenneth J. D. Allen, AM Department of Psychology Harvard University Ananda B. Amstadter, Amstadter, PhD Virginia Institute for Psychiatric and Behavioral Genetics Virginia Commonwealth University Michael F. Armey, PhD Department of Psychiatry and
Department of Psychology Emory University April L. Brown, Brown, MPH Department of Psychology Emory University Mindy Brown, BS Department of Psychology Te University of Utah Alexander L. Chapman, PhD, RPsych RPsych Department of Psychology Simon Fraser University Dante Cicchetti, PhD
Human Behavior W Warren arren Alpert Medical School Brown University Rachel M. Atchley Atchley,, PhD, MCR Center on Mindfulness and Integrative Health Intervention Development (C-MIIND) Te University of Utah Teodore P. Beauchaine, PhD Department of Psychology Te Ohio State University Lane Beckes, PhD College of Liberal Arts and Sciences Bradley University Spencer Bell, PhD Department of Physiology/Pharmacology Wake W ake Forest University Health Health Sciences Ziv E. Bell, MA Department of Psychology Te Ohio State University Michele Berk, PhD Psychiatry and Behavioral Sciences - Child and Adolescent Psychiatry and Child Development Stanford University
Institute Development University Univers ityofofChild Minnesota Pamela M. Cole, PhD Department of Psychology Te Pennsylvania State University Lindsey C. Conkey Conkey,, PhD Department of Psychological and Brain Sciences University Univers ity of Massachusetts Amherst Elisabeth Conradt, PhD Department of Psychology Te University of Utah Geoffrey W. Corner, Corne r, BS Department of Psychology University Univers ity of Southern California Sheila E. Crowell, PhD Department of Psychology Te University of Utah Katherine L. Dixon-Gordon, PhD Department of Psychological and Brain Sciences University Univers ity of Massachusetts Amherst Anna R. Docherty, Docherty, PhD University Univers ity Neuropsychiatric Institute University Univers ity of Utah Health Sciences Center
Grace Binion, MS Department of Psychology University Univers ity of Oregon
Weston W eston Layne Edwards, MA Department of Psychology Bradley University
S H O RT C O N T E N T S
About the Editors Contributors
ix
Table of Contents Chapters Index
vii
xiii
1–486
487
A B O U T T H E E D I TO R S
psychology ology Teodore P. P. Beauchaine, Beauch aine, PhD, Ph D, earned his undergraduate degree in psych from Portland State University, University, and his PhD in clinical cl inical psychology, psychology, with a quantitative minor, from Stony Brook University. University. He completed his clinical internship at the University of California at San Diego School of Medicine. He is past recipient of both the American Psychological Association Distinguished Scientific Award Awar d for Early Career Contributions to Psychology and the American Psychological Association Mid-Career Award for Outstanding Contributions to Benefit Children, Youth, and Families. He has served on numerous editorial boards, and as Associate Editor for Development and Psychopathology and Psychophysiology.. He served on the National Institute of Mental Health National Psychophysiology Advisory Council Workgroup on asks asks and Measures for the Research Domain Criteria (RDoC). His research addresses neural underpinnings of and development of behavioral impulsivity impulsivit y, emotion dysregulation, and intentional intenti onal self-injury in children, adolescents, and adults. Sheila E. Crowell earned her PhD in child clinical psychology from the
University of Washington. She completed her clinical internship at Seattle Children’s Hospital through the University of Washington Psychology Internship Program. Dr. Crowell has expertise in emotion dysregulation across the lifespan, including infants, children, adolescents, and adults. Her work on emotion dysregulation extends across a number of diverse clinical cli nical populations, such as depression, substance use disorders, trauma, personality disorders, and self-injury. Dr. Crowell is also a licensed clinical psychologists with expertise in Dialectical Behavior Terapy (DB), an evidence-based treatment for diagnoses characterized by emotion dysregulation. Dr. Crowell has served on study sections for the National Institutes of Health and as a reviewer or editorial board member for several journals. She has received funding for her research from the National Institutes of Mental Health and the American A merican Foundation for Suicide Prevention. A primary goal of Dr Dr.. Crowell’ Crowell’ss research is to prevent suicide and the development development of psychopathology through enhanced identification of those at risk and early intervention.
Margaret A. Fields-Olivieri Department of Psychology Te Pennsylvania State University Courtney N. Forbes, MEd Department of Psychology Te University of oledo Brett Froeliger, PhD Department of Neuroscience Medical University of South Carolina Eric L. Garland, PhD, LCSW Center on Mindfulness and Integrative Health Intervention Development (C-MIIND) Te University of Utah Kim L. Gratz, PhD Department of Psychology Te University of oledo James J. Gross, PhD PhD Department of Psychology Stanford University Hunter Hahn, MA Department of Psychology Te Ohio State University Nathaniel Haines, BA Te Center for Cognitive and Brain Sciences Te Ohio State University Greg Hajcak, PhD Department of Psychology Florida State Univ University ersity Lauren A. Haliczer, MA Department of Psychological and Brain Sciences University Univers ity of Massachusetts Amherst Sophie Havighurst, PhD
Hooria Jazaieri, PhD Kellogg School of Management Northwestern University Parisa R. Kaliush, BA Department of Psychology Te University of Utah Niranjan S. Karnik, MD, PhD Department of Psychiatry Rush Medical College Erin A. Kaufman, BA Department of Psychology Te University of Utah Christiane Kehoe, PhD Department of Psychiatry Te University of Melbourne Patricia K. Kerig, PhD Department of Psychology Te University of Utah Mona Khaled, MA Department of Psychology University Univers ity of Southern California Joseph C. Leshin, BS Department of Psychology & Neuroscience Te University of North Carolina at Chapel Hill Kristen A. Lindquist, PhD Department of Psychology & Neuroscience Te University of North Carolina at Chapel Hill Christina Gamache Martin, PhD Department of Psychology University Univers ity of Oregon Whitney I. Mattson, PhD Brain Development and Social
Department of of Psychiatry Te University Melbourne Sage E. Hawn, BA Virginia Institute for Psychiatric and Behavioral Genetics Virginia Commonwealth University Nora H. Hope, PhD, RPsy R Psych ch Department of Psychology Simon Fraser University Sarah A. Horvath, MA Department of Psychology Ohio University Camelia E. Hostinar Hostinar,, PhD Department of Psychology University Univers ity of California, Davis
Cognition Lab Nationwide Children’s Hospital Kateri McRae, PhD Department of Psychology University Univers ity of Denver Michele A. Morningstar Morningstar,, PhD Brain Development and Social Cognition Lab Nationwide Children’s Hospital Eric E. Nelson, PhD Center for Biobehavioral Health Nationwide Children’s Hospital Emily Neuhaus, PhD Department of Psychology University of Washington
Jacqueline O’Brien, MS Department of Psychology University Univers ity of Oregon Cassie Overstreet, MA Department of Psychology Virginia Commonwealth University Ruchika Shaurya Prakash, PhD Department of Psychology Te Ohio State University Sarah E. Racine, PhD Department of Psychology McGill University K. Ashana Ramsook, MA Department of Psychology Te Pennsylvania State University Lance M. Rappaport, PhD Virginia Institute for Psychiatric and Behavioral Genetics Virginia Commonwealth University Julia R. Richmond, MA Department of Psychology Te University of oledo
Ross A. Tompson, PhD Department of Psychology University Univers ity of California, Davis Matthew . ull, PhD Department of Psychology Te University of oledo Andero Uusberg, PhD Department of Psychology University of artu artu Helen Uusberg, PhD Department of Psychology University of artu artu Robert D. Vlisides-Henry Vlisi des-Henry,, BA Department of Psychology Te University of Utah Gemma . Wallace, BA Department of Psychiatry University Univers ity of Utah School of Medicine Sara F. Waters Waters,, PhD Department of Human Development Washington W ashington State University University,,
Darby Saxbe, PhD Department of Psychology University Univers ity of Southern California Heather . Schatten, PhD Department of Psychiatry and Human Behavior Brown University iffany M. Shader, MA Department of Psychology Te Ohio State University Brittany C. Speed, MA Department of Psychology
Vancouver Linnie E. Wheeless, JD Department of Psychology Te University of oledo Patrick Whitmoyer, MA Department of Psychology Te Ohio State University Dominika A. Winiarski, PhD Department of Psychology Rush University Maureen Zalewski, PhD Department of Psychology
Stony University Sarah A. Brook Stoycos, MA Department of Psychology University Univers ity of Southern California
University ity ofPhD Oregon ParUnivers Paree ee Zarolia, Department of Psychology Te University of Denver
TA B L E O F C O N T E N T S
. Functionalist and Constructionist Perspectives on Emotion
. . . .
. .
. .
.
Dysregulation Teodore P. Beauchaine and Nathaniel Haines Emotions as Regulators of Motivated Behavior Eric E. Nelson, Michele A. Morningstar, Morningstar, and Whitney I. Mattson Emotions as Regulators of Social Behavior Lane Beckes and Weston Layne Edwards Cognition and Emotion in Emotion Dysregulation Kateri McRae and Paree Paree Zarolia What Emotion Dysregulation Looks Like: Inferences Inferences from Behavioral Observations K. Ashana Ramsook, Pamela M. Cole, and Margaret A. Fields-Olivieri Emotion Dysregulation and Aging
Patrick Whitmoyer and Ruchika Shaurya Prakash Emotion Generation, Generation, Regulation, and Dysregulation as Multilevel Transdi ransdiagnosti agnosticc Constru Constructs cts Sheila E. Crowell, Robert D. Vlisides-Henry, and Parisa R. Kaliush Development of Emotion Dysregulati Dysregulation on in Developing Relationships Ross A. Tompson and Sara F. Waters Operant Reinforcement and Development of Emotion Dysregulation Christina Gamache Martin, Maureen Zalewski, Gr Grace ace Binion, and Jacqueline O’Brien O’Brien Cognitive Processes and Risk for Emotion Dysregulation Hooria Jazaieri, Helen Uusberg, Andero Uusberg, and James J. Gross
. Interpersonal Processes and the Development Development of Emotion DysregulationProcesses Sarah A. Stoycos, Geoffrey W. Corner, Mona Khaled, and Darby Saxbe . Respiratory Sinus Arrhythmia as a Transdiagnostic Transdiagnostic Biomarker of Emotion Dysregulation Teodore P. Beauchaine and Ziv E. Bell . Event-Related Potentials and Emotion Dysregulation Brittany C. Speed and Greg Hajcak . Neuroimaging of Emotion Dysregulation Joseph C. Leshin and Kristen A. Lindquist . Behavioral and Molecular Genetics of Emotion Dysregulation Lance M. Rappaport, Sage E. Hawn, Cassie Overstreet, and Ananda B. Amstadter Amstadter
. Epigenetic Foundat Foundations ions of Emotion Dysregulati Dysregulation on Mindy Brown, Elisabeth Elisabeth Conradt, and Sheila Sheila E. Crowell Crowell . Emotion Dysregulatio Dysregulation n and Externaliz Externalizing ing Spectrum Disorders iffany M. Shader and Teodore P. Beauchaine . Emotion Dysregulatio Dysregulation n and Internalizi Internalizing ng Spectrum Disorders Camelia E. Hostinar and Dante Cicchetti . Emotion Dysregulatio Dysregulation n and Childhood rauma Patricia Pa tricia K. Kerig . Emotion Dysregulatio Dysregulation n in Autism Spectrum Disorder Emily Neuhaus . Emotion Dysregulatio Dysregulation n and Psychosis Spectrum Disorders Gemma . Wallace and Anna R. Docherty . . Emotion Dysregulatio Dysregulation n in Addiction Eric L. Garland, Spencer Bell, Rachel M. Atchley Atchley,, and Brett Froeliger . Emotion Dysregulatio Dysregulation n and Eating Disorders Sarah E. Racine and Sarah A. Horvath . . Emotion Dysregulatio Dysregulation n and Self-Inflicted Injury Erin A. Kaufman and Sheila E. Crowell . Emotion Dysregulatio Dysregulation n and Borderline Personality Disorder Katherine L. Dixon-Gordon, Lauren A. Haliczer Haliczer,, and Lindsey C. Conkey . . Behavioral Assessment of Emotion Dysregulat Dysregulation ion Molly Adrian and Michele Berk Berk . Self-Report Assessme Assessment nt of Emotion Dysregulation Kim L. Gratz, Courtney N. Forbes, Linnie E. Wheeless, Julia R. Richmond, and Matthew . ull ull . . Assessme Assessment nt of Emotion Dysregulation Using Ecological Momentary Assessment Heather . Schatten, Kenneth J. D. Allen, and Michael F. Armey . reating Emotion Dysregulation in Externaliz Externalizing ing Disorders Dominika A. Winiarski, April L. Brown, Niranjan S. Karnik, and Patricia A. Brennan . . reating Emotion Dysregulation in Internalizin Internalizingg Disorders Christiane Kehoe and Sophie Havighurst . Dialectical Behavior Terapy and reatment reatment of Emotion Dysregulation Alexander L. Chapman and Nora Nora H. Hope Hope . Future Directions in Research Rese arch and reatment reatment of Emotion Em otion Dysregulation Teodore P. Beauchaine, Hunter Hahn, and Sheila E. Crowell Index
C H A P T E R
Functionalist and Constructionist Functionalist Perspectives on Emotion Dysregulation
Theodore P. Beauchaine and Nathaniel Nathaniel Haines
Abstract
Two theoretical perspectives—functionalism and constructionism—predominate modern research on emotion. This introductory chapter describes these perspectives and offers points of convergence and divergence. It pays special attention to common misconceptions about functionalism and to strengths and limitations of each perspective. Functionalism, which draws in part from phylogenetic accounts of emotion and motivation, is limited by difficulties drawing inferences about human emotion from animal research, even though animal research is conducted using very precise methods of high spatial and temporal resolution. In contrast, constructionism is limited by difficulties falsifying its core propositions given reliance on research using functional magnetic resonance imaging, which has poor temporal resolution.. These limitations notwithstanding, both functionalism and constructionism have much to resolution offer current interpretations of and future research on emotion dysregulation. Thus, pitting the perspectives against one other is counterproductive. Keywords: constructionism constructionism,, emotion, emotion dysregul dysregulation, ation, functionalism, psychop psychopathology athology
Work on this chapter Work chapter was supported supported by grant grant DE DE from the the National National Institutes of Health, and by the Nationa Nationall Institutes of Health Science of Behavior Change (SoBC) Common Fund. Introduction
It is an honor a privilege to coedit this diverse Oxford Handbook , in and which contributors describe perspectives on emotion dysregulation. We were fortunate to receive contributions from internationally renowned experts in affective science, who together summarize contemporary approaches to and future directions in emotion dysregulation research. Chapters are grouped into six sections: () conceptual issues; () cognitive, behavioral, and social approaches; () neurobiological approaches; () psychopathology; () assessment and treatment; and () future directions. Collectively, Collectively, these sections describe effects of emotion dysregulation on core aspects of human function across levels of analysis including genes, neural networks, electrophysiology, and behavior. During years of planning and
editing this volume, Sheila Crowell and I (TPB) learned more about than we otherwise could haveemotion known,dysregulation and we are indebted to a brilliant team of contributors. We hope readers find the diversity of topics useful in advancing their thinking about emotion dysregulation and its multiple determinants across the lifespan. In this chapter,, we summarize functionalist and constructionist ter perspectives on emotion, which sets the stage for chapters to follow. Over the past two decades, emotion regulation regulation has received burgeoning attention as a scientific construct, as evidenced by foundational articles, dedicated volumes, and extended scientific debate (e.g., Aldao, Nolen-Hoeksema, & Schweizer, ; Cole, Martin, & Dennis, ; ; Gross, , ). Although emotion dysregulation has received more more
circumscribed attention, it is of considerable inter-
One common use of the term functionalism functionalism as-
est to developmentalists, psychopathologists, and other invested parties (e.g., Beauchaine, ; Bradley et al., ; Gratz, Rosenthal, ull, Lejuez, & Gunderson, ; Linehan, ). In this volume, we plac placee prima primary ry empha emphasis sis on emot emotion ion dysr dysregul egulation ation and how it compromises adaptive human functioning through its effects on initiating, maintaining, and modulating diverse human behaviors (cf. Campos, Mumme, Kermoian, & Campos, ; Tompson, ). Given our objective of conveying contemporary perspectives on emotion dysregulation, both emotion and emotion regulation must be discussed. However, they are not primary foci given widespread coverage in other sources. Interested readers are referred to excellent recent reviews (Aldao et al., ; Barrett, a; Braunstein, Gross, & Ochsner, ; Gross, ; Gross & Barrett, ).
When defining emotion dysregulation, one must first consider what emotions are, and the day-to-day functions they serve and do not serve in both their ordinary and extreme forms. From this perspective,
sumes evolutionary selection of at least some human emotions. Such accounts presume that broad classes of emotion evolved to motivate adaptive, survivalrelated functions including approach, avoidance, and social affiliation (e.g., Keltner & Gross, ). According Accor ding to evolutio evolutionary nary functiona functionalist list perspec perspectives, tives, emotions that subserve these functions are preserved across species and experienced by all mammals, including humans, because they were selected in our environments of adaptation (e.g., Panksepp, , ). For example, approach emotions (e.g., wanting, enthusiasm) elicit consummatory behaviors (e.g., foraging, food seeking); avoidance emotions (e.g., anxiety, fear) elicit precaution (e.g., passive avoidance, suppression of approach); and affiliative emotions (e.g., compassion, affection) elicit prosocial behaviors (e.g., group cohesion, pair bonding). Without Witho ut emotions emotions motivating motivating approach, approach, avoidance avoidance,, and affiliative behaviors, likelihood of survival in our environments of adaptation would presumably have been lower. Evolutionary functionalist perspectives have a long history in animal, human, and comparative research on emotion and suggest that
affect dysregulation cannot be defined by overt expressions of emotion without first specifying the contexts in which such expressions occur, then evaluating whether the emotion expressed and the intensity of its expression are context appropriate, inappropriate,, or neutral vis-à-vis social and cultural inappropriate norms. For example, intense expressions of anger toward others may be fully functional if the safety of one’s offspring is threatened, but similarly intense displays of anger interfere with adaptive behavior in most social and cultural contexts. Although often not considered, it is also important to note that in some situations expressions of anger are afunctional . Even moderately intense solitary displays of anger, for example, such as those elicited by frustration while driving, driving, may may serve no function function or dysfunction whatsoever.. Tus, whether particular displays of whatsoever emotion are functional, dysfunctional, or afunctional, and whether they are regulated, dysregulated, or unregulated, depends in large part on eliciting contextual events, and match or mismatch between context and expressive intensity (e.g., Aldao, ). Furthermore, given two common uses of the term functionalism functionalism that that partly but do not fully overlap (see immediately below), classifying emotions as functional or dysfunctional, regulated or dysregulated, is not as straightforward as it might first appear (e.g., Barrett, b).
emotion and motivation are inextricable facets of human function, despite being separated in the history of behavioral science (see, e.g., Beauchaine & Zisner, ; Gray & McNaughton, ; Panksepp, ; Porges, ). An important corollar corollaryy of this perspective is that humans sometimes behave at the behest of their emotions. Such is especially likely when environmental contingencies are extreme and pull strongly for survival-relevant actions (e.g., in situations of food deprivation, threats to physical safety to oneself or one’s kin). Strong emotional reactions to these situations motivate urgent behavioral responses that override ongoing activities (see, e.g., Corr, ). Notably, however, evolutionary functionalist accounts do not imply that all or even most emotional reactions are survival relevant. In fact, evolutionary theorists have long recognized that () over any extended period of time individual differences in emotional and behavioral response tendencies confer probabilistic rather than deterministic effects on adaptive fitness, and () some behavioral response tendencies are coincidental byproducts of evolution—not direct outcomes of adaptive selection (Beauchaine, ; Buss, Haselton, Shackelford, Bleske, & Wakefield, W akefield, ; Gould, ). In the latter case, such response tendencies have no bearing on
Variants V ariants of Functionalism
adaptive fitness. Despite its name, evolutionary
circuitry is a difficult proposition (see Barrett, b).
functionalism therefore therefore does not imply that all or even most emotional experiences or expressions are functional, a point we return to in later sections (see “Points of Divergence and Convergence in Functionalism and Constructionism”). In a second common use of the term functi term functionalonalism,, experiences and expressions of emotions are ism linked to outcomes in our day-to-day lives, with limited if any consideration of our evolutionary environments of adaptation (see Keltner & Gross, ). Among children, for example, emotionally complaisant, well-mannered behavior in the classroom is seen as functional and adaptive, whereas emotionally exuberant, impulsive behavior is seen as dysfunctional and maladaptive. Notably, however, exuberance and impulsivity were likely not maladaptive in our evolutionary environments of adaptation and may have conferred selective advantages in certain environmental niches (see Mead, Beauchaine, & Shannon, ). Tus, whether specific emotions and behaviors are functional or dysfunctional in our modern-day lives may have nothing to do with their phylogenetic adaptive value. Our intent here is to
Tese observations contributed to modern constructionist accounts of emotion, which eschew several assumptions of traditional functionalist theories, as described in the next section. A common although not universal assumption of evolutionary functionalism is that at least some emotions or subsets of emotions represent categories in nature. Tis notion follows from seemingly different classes of behavior—including approach, avoidance, and social affiliation—that specific emotions seem to support (see earlier; Beauchaine & Zisner, ; Gray Gray & McNaughton, ; Panksepp, Pankse pp, ; Panksepp & Watt, ). Furthermore, many evolutionary accounts presume that either rudiments of or fully formed approach, avoidance, and affiliative emotions () are present across mammalian species, () are experienced by human infants at birth, and () transcend human cultures (Ekman & Cordaro, ; Ekman & Friesen, ). Tese theories articulate phylogenetically old, subcortical neural networks that subserve basic emotions (see, e.g., Beauchaine, Neuhaus, Zalewski, Crowell, & Potapova, ; Panksepp, ).
call readers’ attention to the important distinction between these two common uses of the term functerm functionalism,, the latter of which is especially prone to tionalism circular reasoning in definitions of adaptation. Given potential confusion brought about by different uses of the term functionalism term functionalism,, and given other issues described in foundational articles across the affective sciences (e.g., Campos et al., ; Cole et al., ; Keltner & Gross, ; Tompson, ), coeditor Sheila Crowell and I (PB) ( PB) encouraged authors to adopt a common definition of emotion dysregulation as “a pattern of emotional experience and/or expression that interferes with appropriate goal-directed behavior ” (Beauchaine, , p. , emphasis added; see also Cole, Hall, & Hajal, ). Here, we chose the word appropriate instead of adaptive to to avoid teleological undertones. eleological explanations are those that define phenomena based on specific purposes they serve, including assumptions that specific emotions either () evolved to serve highly specialized functions or () always serve an immediate function. As already noted, many displays of emotion are afunctional, and in Western culture, situations that require “appropriate” dampening of strong emotions are far removed from our evolutionary environments of adaptation. Moreover, testing evolutionary func-
Full articulation of anatomical and functional characteristics of these subcortical structures is beyond the scope of this introductory chapter, but both are specified in extensive reviews of the animal and human literatures (e.g., Beauchaine et al., ; Ikemoto, Yang, & an, an, ; ; Koob & Volkow, ; Panksepp, ; ovote, Fadok, & Lüthi, ). In brief, early work on subcortical neural circuits of approach and avoidance derived from studies of associative learning, motivation, and addiction in rodents and nonhuman primates. Tis work, including lesion studies, single cell recording experiments, and pharmacological manipulations, identified subcortical neural systems of appetitive and aversive motivation that are largely preserved across species. Tese systems include () the medial forebrain bundle—particularly projections from the ventral tegmental area to the nucleus accumbens— as integral to appetitive motivation and approach emotions (Sagvolden Johansen, Aase, & Russell, ; Schultz, ; Wise, Wise, ) and () the septosepto hippocampal system—including the hippocampus and its afferent projections from the amygdala—as integral to aversive motivation and associated avoidance emotions (Corr, ; Gray & McNaughton, ; Strange, Witter, Lein, & Moser, ). Although general consensus exists regarding the
tions of emotions and emotion-subserving neural
primary roles these systems play in approach and
avoidance motivation and emotion (for reviews see
; Galvan et al., ). Differential neuromatuneuromatu-
Beauchaine, ; Beauchaine & Zisner, ), it is also well recognized that the systems interact structurally and functionally (e.g., Corr, ; Corr & McNaughton, ). For example, neurons in the nucleus accumbens (NAcc) respond to punishment as well as reward, and the amygdala responds to reward as well as punishment (e.g., Sauder, Derbidge, & Beauchaine, ; Schultz, ). Both are therefore intricately involved in associative learning. Moreover, the NAcc and the amygdala share inter-
ration of subcortical and cortical brain regions is a likely contributor to the impetuous, impulsive, and emotionally labile behaviors common to adolescence (e.g., Casey & Caudle, ). As prefrontal neuromaturation completes in early adulthood, self- and emotion regulation improve markedly. Notably, children and adolescents show stronger subcortical responses to incentives than adults, yet their prefrontal cortex responding is weaker and more diffuse (Macdonald, Goines, Novacek, &
connections via the paraventricular nucleus and the stria terminalis (e.g., Dong, Li, & Kirouac, ; ovote et al., ). Tus, although the distinction between appetitive and aversive subcortical systems is useful heuristically heuristically,, subcortical CNS networks of approach and avoidance interact complexly and are not functionally independent (see also Beauchaine & Constantino, ; Beyeler, ).
Walker,, ). Furthermor Walker Furthermore, e, adolescents with impulse control problems show blunted frontal neuromaturation (De Brito et al., ). Finally, deficits in functional connectivity between cortical and subcortical structures are observed in both impulse control and anxiety disorders, which are characterized by excessive approach- and avoidance-related emotions, respectively. For example, reduced functional connectivity between the anterior cingulate and dorsal striatum is observed among externalizing adolescents (e.g., Shannon, Sauder, Beauchaine, & GatzkeKopp, ), and reduced functional connectivity
Implications for Emotion Regulation and Dysregulation Functionalists often distinguish between bottomup, subcortically mediated emotion generation processes and processes top-down,(e.g., cortically mediated emotion regulation Beauchaine, ; Gross & Barrett, ). According to such perspectives, subcortical neural circuits that initiate strong emotional responses are modulated by cortical functions (see also Hare et al., ). Tis literature is voluminous and cannot be reviewed comprehensively, yet several findings are noteworthy. First, cortical structures, particularly in prefrontal and orbitofrontal regions, have long been implicated in executive function and self-regulation (see Beauchaine & Zisner, ; Etkin, Büchel, & Gross, ; Heatherton, ). Modern neuroimaging studies identify functional subdivisions of the prefrontal, anterior cingulate, and insular cortices as integral to effortful downregulation of negative affect (e.g., one, Garn, & Pine, ; Zilverstand, Parvaz, & Goldstein, ). In fact, volitional reappraisal of negative emotion elicits increased neural responding across a distributed network of frontal structures, including the dorsolateral, medial, and ventrolateral prefrontal cortices; the lateral orbitofrontal cortex; the inferior frontal gyrus (IFG); and the insular cortex (e.g., Goldin, McRae, Ramel, & Gross, ). Second, subcortical structures reach volumetric and functional maturity many years before cortical
An alternative to functionalist perspectives is constructionist theory. Constructionists assert that what humans perceive as discrete emotions are not encoded by specific brain regions, but rather constructed through learning processes that are highly individualized. According to this perspective, emotions and other experiential states, including perception and cognition, emerge from interactions
neural structures (e.g., Brain Development Cooperative Group, ; Casey, Getz, & Galvan,
among more primitive sensory and neural mechanisms, which humans interpret and categorize
between the amygdala and the orbitofrontal is associated with compulsive behavior andcortex emotional lability (e.g., Churchwell, Morris, Heurtelou, & Kesner, ; Hilt, Hanson, & Pollak, ). Notably, although findings are complex and not fully consistent, several studies show improved cortical–subcortical connectivity following effective treatment for internalizing and externalizing disorders (for a recent review see Beauchaine, Zisner, & Hayden, ). Collectively, these findings lend support to the notion that emotion regulation is subserved by top-down cortical control over subcortical neural responding, and that disruptions in frontal cortical function and cortical–subcortical connectivity characterize emotion dysregulation (see also Beauchaine, Constantino, & Hayden, ).
Constructionism
based on prior experience (see Barrett, ). Constructionist theory identifies core affective pro-
construed in critical discussions concerning its merits. Tis creates an artificial distinction between
cesses , including valence and arousal , which transcend multiple emotional states. Trough repeated visceral pairings of these core affective processes with sensory and neural input elicited by our environments, we learn to associate instances of core affect with higher order, discrete representations of emotion such as happiness and sadness (e.g., Russell & Barrett, ). Neural mechanisms of core affective states are presumed to be present at birth, uni-
functionalist and constructionist views on emotion. Wee view functionalism and constructionism as W largely compatible, so long as one avoids teleological misconceptions of evolution and acknowledges interactive complexities and functional dependencies of neural responding within and across subcortical and cortical networks. In sections to follow, we briefly outline our reasoning.
versal among humans, and supported by the same neural networks as other psychological processes and states, such as perception and decision making (see Duncan & Barrett Barrett,, ). Constructionists make a clear distinction between core affective processes and emotions. Whereas core affective processes refer to general experiences of positivity–negativity (valence) and activation–deactivation (arousal), emotions are more specific experiential states, such as sadness, anger, fear, and shame (Ekman, ; Ekman & Cordaro, ). Tus, despite being experienced discretely, all emotions can be characterized along dimensions of
Implications for Emotion Dysregulation
Constructionist accounts now rival functionalist perspectives as explanatory theories of emotion, yet constructionists have written far less than functionalists about emotion dysregulation per se. Tis may be because constructionist approaches, including the theory of constructed emotion (CE; Barrett, a, b; Lindquist, ), view emotions as emergent properties of complex neuro-architectures, which exhibit individualized affect-imbuing response patterns that are byproducts of unique learning histories. Tese learning histories produce cognitive–affective schemas, attributional biases,
valence and arousal (Barrett, ). to constructionist theory, we rely on According learning and memory from prior experience to infer the meaning of core affect in current situations. In this way, we construct context-dependent emotion representations (Barrett, a). Of note, core affective processes, similar to basic emotions, can motivate behavioral response tendencies. tenden cies. For example, Pavlovian bias is a “hard-wired” tendency to approach positively valenced stimuli and avoid negatively valenced stimuli (Guitart-Masip et al., ). In some cases, Pavlovian bias is so strong that organisms cannot learn stimulus–response contingencies that require avoidance to attain reward (Hershberger, ). Constructionists have been critical of functionalist theories on a number of grounds. Although we cannot review all such critiques here, three especially important issues concern () teleological arguments concerning adaptive evolution of emotion, as outlined earlier (see “Variants of Functionalism”); () opposition to the notion that specific neural structures and networks subserve particular emotional states and functions; and () disagreement on the extent to which animal research on reward learning, fear learning, and motivation informs our understanding of human emotion. Although we agree that these points warrant consideration when
and stimulus–response associations that From contribute collectively to emotional experience. this standpoint, emotion dysregulation is emotion, emotion, because it arises through the same highly individualized neural processes and unique learning histories (see, e.g., Papa & Epstein, ). According to CE, emotion dysregulation emerges at least in part from neural mechanisms of core affective processes (e.g., valence, arousal) and disruptions in situated conceptualization conceptualization (Barrett, Wilson-Mendenhall, Wilson-M endenhall, & Barsalou, ). Situated conceptualization conceptualizatio n refers to “the brain [as] a situated processing architecture, designed to process situations in the moment and to simulate non-present situations in thought” (Barsalou, , p. ). Tis includes evaluating what eliciting events represent, how to act upon those events, and the nature of core affective processes to expect. Barrett et al. () suggest that emotion dysregulation could emerge from highly canalized, inflexible conceptualizations that are not situational. In turn, nonsituational conceptualizations could arise from disruptions to one or more among several processes, including memory retrieval, autonomic regulation, and attention, to name but a few. As reviewed by Barrett and Satpute (), many such deficits correlate with abnormalities in connectivity among intrinsic neural net-
evaluating functionalist theories, we argue that functionalism is often oversimplified and thus mis-
works, including the salience network and the fronto-parietal network, as seen in diverse forms of
psychopathology. Tus, disrupted connectivity plays a central role in both functionalist and con-
A second critique of functionalism concerns its linking of emotional states to specific brain
structionist theories of emotion dysregulation.
As already already noted, teleological explanations are those those that define phenomena based on the specific purposes they serve. Te notion that emotions were designed by evolution to serve specific, adaptive functions is therefore teleological (Barrett, b). Basic emo-
regions and networks. As outlined under “Variants of Functionalism,” for example, functionalist theories often link () appetitive emotions to neural responding in the medial forebrain bundle, including projections from the ventral tegmental area to the nucleus accumbens (Sagvolden et al., ; Schultz, ; Wise, ), and () aversive emotions to neural responding in the septo-hippocampal system, including the hippocampus and its afferent pro-
tion theory identifies approximately six discrete emotions (happiness, sadness, fear, surprise, anger, and disgust) that are shared across cultures, some of which are documented in other mammals (Chevalier-Skolnikoff, ; Ekman & Friesen, ). A teleological explanation takes cross-species and cross-cultural expressions of affect as evidence that discrete emotions evolved to facilitate adaptive behaviors (e.g., fear evolved with the purpose of signaling organisms to danger). In other words, evolution by natural selection purposefully designed basic basic emotional states to preserve the organism. Here it is important to note that purposeful
jections from the amygdala (Corr (Corr,, ; Gray & McNaughton, ; Strange et al., ). According to such accounts, basic emotions are presumed to be initiated/generated by localized, phylogenetically old neural structures that are largely preserved across mammals and, in some cases, other vertebrates (Panksepp, , ). As already noted, these conclusions are based on decades of extensive research with animals (rodents and nonhuman primates). Tis research includes localized lesion studies, pharmacological manipulations, and implanted electrode stimulation and recording experiments that are highly precise both spatially and temporally
design has been explicitly eschewed a mechanism On the of evolution since Darwin () aswrote Origin of Species . Tus, even though evolutionary psychologists have at times appealed to purposeful design, evolutionary biologists rejected the notion over a century ago (see Beauchaine, ; Buss et al., ; Gould, ). A more accepted approach among evolutionary theorists is to infer function from the consequences of emotions throughout evolutionary history (Wright, ). For example, we may infer that the function of fear is to alert an organism of immediate danger because fear in the face of danger creates conditions that are conducive to survival. By way of natural selection, the most likely consequence of an emotion is therefore the function of that emotion specifically in our environments of evolutionary adaptation (see Wright, ). Provided Pro vided such inferences are supported by observable, biological mechanisms, they are not dubious philosophically (e.g., Barrett, b; Neander, ). Tis Darwinian () perspective is infused in contemporary thinking about emotion (see Keltner & Gross, ), despite terminology that sometimes leads to confusion. Indeed, even in biology, where most scientists decry language implying that evolution proceeds with goal-directed intention, some nevertheless use such language as a literary
(e.g., Gray, ;ofOlds Milner, ). Most studies this & nature cannot be conducted with humans for obvious ethical reasons. As a consequence, neural studies of human emotion rely primarily on functional magnetic resonance imaging (fMRI), which evolved more recently. When fMRI technology was first applied in emotion research, region-of-interest (ROI) and effective connectivity analyses predominated. Most early ROI and connectivity studies were deductive (top-down), with ROIs specified a priori based on theory. Early metaanalyses of these studies revealed modest evidence for emotion localization, consistent with animal research (e.g., Murphy, Nimmo-Smith, & Lawrence, ; Phan, Wager, Wager, aylor, aylor, & Liberzon, Liber zon, ). ). More recently, inductive (bottom-up) approaches that capture coactivated neural circuitry have ascended to prominence in the fMRI literature. Tese approaches show that distributed patterns of neural activity account for more variance in basic emotions than specific brain regions (Celeghin, Diano, Bagnis, Viola, & amietto, ; Saarimäki et al., ). Such findings are sometimes cited as evidence against functionalism (Kober et al., ; Lindquist, Wager, Kober, Bliss-Moreau, & Barrett, ; ouroutoglou, Lindquist, Dickerson, & Barrett, ). It is im-
device (e.g., Hanke, ).
portant to note, however, that linking basic
Points of Divergence Divergence and Convergence in Functionalism and Constructionism
eemotions motions to specific brain structures oversimplifies the functionalist perspective. In fact, functionalists
In contrast, functionalist accounts suggest that regulated emotions should be associated with ()
have long recognized that multiple emotional states activate common brain regions (see, e.g., Gray & McNaughton, ), and that complex neural circuits and interacting cortical–subcortical networks generate and regulate affective responses (e.g., Beauchaine, ; Beauchaine & Zisner, ; Braunstein et al., ; Etkin et al., ; Goldin et al., ; Gray & McNaughton, ; Gross & Barrett, ). Indeed, functional defi-
subcortically generated responses to eliciting events that are () modulated by cortical responses via () strong cortical–subcortical connectivity (e.g., Beauchaine, Constantino, et al., ; Beauchaine, Zisner, et al., ). If such is the case, we would expect to find distributed neural activity for any given instance of emotion due to the limited temporal resolution of fMRI. From this perspective, prominent functionalist accounts look much like
ciencies in cortical–subcortical connectivity characterize several psychiatric disorders for which emotion dysregulation plays a prominent role (see, e.g., Beauchaine, Constantino, Constantino, et al., ; Shannon Shanno n et al., ; one et al., ). It should also be noted that neural signals propagate across brain regions and networks at much faster rates than fMRI is capable of resolving. For example, reactivity to reward cues by midbrain dopamine neurons—as assessed via direct electrode placement in primates—peaks at about . seconds and returns to baseline at about . seconds (e.g., Lak, Stauffer, & Schultz, ). In contrast,
contructionist ones—both assume that primaryprocess emotions (core (core affective states ) that promote approach and avoidance behaviors are subserved by phylogenetically old structures (primarily subcortical structures), and that cortical networks interact with these subcorti subcortical cal networks to produce what we consciously experience as emotional states (see Panksepp, ). Finally, the locationist basic basic emotion perspective of functionalism is only a single perspective—albeit a pervasive one in certain areas of research. Other functionalist perspectives focus on the dimensional nature of emotions and various contextual
the fMRI blood oxygen level–dependent (BOLD) signal peaks between and seconds poststimulus and returns to baseline after seconds (e.g., Lohrenz, Kishida, & Montague, ). Tus, the BOLD signal is a sluggish indicator of neural responding and is not well suited for detecting rapidly propagating patterns of neural responding that originate in the subcortex and project to both cortical and other subcortical structures. Although modern imaging sequences provide whole-brain coverage of slices at well under -second resolution (e.g., Uğurbil et al., ), this does not circumvent sluggishness of the BOLD signal being measured. It is important to recognize this limitation when evaluating strengths and weaknesses of modern imaging techniques that characterize and correlate widely distributed patterns of BOLD coactivation with attentional and emotional processes (e.g., Yoo et al., ). It remains quite possible that among humans, at least some emotional states arise from patterns of neural reactivity that originate in the very subcortical structures identified in decades of animal research. As described earlier, several functionalist accounts suggest that vulnerability to emotion dysregulation occurs when rapid subcortical responses to eliciting events are not modulated effectively by cortical reactivity (e.g., Beauchaine, ; Casey
factors influence our experience of affect Camposthat et al., ; Haines et al., 2019), similar(cf. to constructivist ideas. In sum, although language used to describe functionalist and constructivist theories is quite different, underlying ideas are more similar than some recent discourse in the literature suggests. A final critique concerns the utility of animal research for making inferences about human emotion. Because evolutionarily functionalism is a phylogenetic account of emotion, many functionalists assume that neural structures implicated in generating basic emotions among humans should overlap considerably (if not fully) with their vertebrate homologues. As noted earlier under “Variants of Functionalism,” such arguments are most persuasive when applied to subcortical regions that are structurally similar across species. Nevertheless, some constructivists have taken a strong stance against comparative research on emotion, noting that functionalist accounts often fail to specify mechanisms adequately, and that it cannot be assumed that emotions are experienced by animals in the same way as they are by humans (see Barrett, b; LeDoux, ). Given () overwhelming structural differentiation of the human cortex, () the phenomenon of human
& Caudle, ; Etkin et al., ).
consciousness, and () the role that language plays in
shaping human experience of emotion, this critique also holds merit. Here again, however, prominent
We look forward We forward to research from both perspectives in upcoming years, and we hope this chapter pro-
functionalists have made the same point. Indeed, Jaak Panksepp, who spearheaded functionalist emotion research with rodents, was very clear in describing qualitative differences between animal and human experiences of emotion: “Are the various affects—diverse feelings of positive and negative valences (‘good’ and ‘bad’ feelings in the vernacular)—identical across species? Of course not! Evolut Evolution ion persistently generates abundant differences, but always on top of conserved-homologous foundational principles at conserved-homologous genetic, neural and primal psychological levels” (Panksepp, , p. ). Our contention is that interspecies differences in emotion notwithstanding, basic animal research offers extensive insights into neural substrates and representations representa tions of emotion, as discussed in previous sections. Models of dopamine reward prediction error signaling in nonhuman primates provide one example (Schultz, Dayan, & Montague, ). Tese models capture moment-to-moment moment-to-moment affective states among humans (Rutledge, Skandali, Dayan, & Dolan, ) and extend further to explain individual differences in mood states, including positive affectivity, irritability, and anhedonia (e.g., Eldar, Rutledge, Dolan, & Niv, ; Laakso et al., ; Zisner & Beauchaine, ). With continued development of computational models of emotion generation and regulation (e.g., Etkin et al., ), we expect that many more such examples will become realized in the near future. Conclusions
In this chapter, we introduce functionalist and constructionist theories of emotion, discuss their implications for understanding emotion dysregulation, and consider points Although of divergence and convergence across perspectives. constructionist theories have gained remarkable traction in affect research and offer key insights into the complex individuality of emotion, we argue that functionalist perspectives still hold value, especially when they are not oversimplified. Functionalist perspectives derive from a long tradition of painstaking neuroscience research, including elegant experiments with animals using techniques of very high spatial and temporal resolution. Although such techniques are not available to those who test constructionist theories with humans, fMRI studies yield insights into the roles that widely distributed neural net works play in emotion and emotion dysregulation.
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C H A P T E R
Emotions as Regulators of Motivated Motivat ed Behavior
Eric E. Nelson, Michele A. Morningstar Morningstar,,
Whitney and Whitney
I. Mattson
Abstract
Emotions, when viewed from the affective neuroscience neuroscience perspective, arise from organized patterns of brain activity, which function to generate adaptive behavioral responses. Behavior that emerges from emotional brain engagement can almost always be characterized as motivated. Thus, emotion and motivation are highly interdependent concepts, particularly when it comes to behavioral expression. However, emotions do not always generate behavior, and behavioral outcomes of emotional engagement—that is, motivated behavior—are not always adaptive. The intersection and dissociation of emotion and motivation are reviewed in this chapter from an affective neuroscience perspective that is heavily influenced by the work of Jaak Panksepp. Keywords: affect, neuroscience, psychopathology, bottom-up, translational
Tis chapter is dedicated to Jaak Panksepp (–), a mentor, scholar, and provocateurr, who brought provocateu brought emotions into the brain brain and the animal animal spirit into humanity.. Jaak was originally slated to write this chapter but his untimely death humanity prevented completion. completion. We have attempted to conjure conjure his intellectual spirit in this writing.
Emotions provide color and meaning to life. Major
attention, facilitate memory, and guide decision making (MacLeod, Mathews, & Tata, ; Cahill
events birth and death are typically bookmarkedsuch withasintense emotional involvement. Even more mundane daily encounters—such as anger directed at an inconsiderate driver, fear of the neighbor’s dog, happiness at seeing one’s family after work—are common punctuations to daily experiences. On a societal level, explorations of emotional experiences are important components of art, literature, and popular culture. All of these elements demonstrate the important role emotion plays in highlighting salient experiences of life. Although these features of of emotion are are clearly of existential importance, biological and psychological perspectives often ascribe a more functional role to emotion. Emotions serve to direct behavior, capture
& McGaugh, Rolls, ; Panksepp Biven, ; Damasio; & Carvalho, ). From & a mechanistic perspective, the similarity and consistency of emotional expression across individuals and species, common neural activation patterns, and similarities in contextual elicitors for a number of emotional experiences suggest that many emotions are conserved across evolution. Furthermore, these expressions serve important roles in fostering life-preserving behaviors, promoting reproductive success, and communicating with conspecifics (Darwin, /; Ekman & Davidson, ; Rolls, ; Panksepp & Biven, ; Damasio & Carvalho, ). Wee view emotion largely from this latter per W spective, and as such approach the topic of emotion
Introduction
more from the standpoint of biological utility than experiential or constructivist perspectives (LeDoux, ; Barrett, ). Core aspects of emotion can be found in fundamental brain response patterns
tions and motivated behaviors evolved because they serve useful biological purposes. However, evolved biological utility is not always adaptive across modern-day contexts, or for all individuals. Next,
(Hamann, ; Panksepp & Biven, ). In both human and animal studies, emotional states such as desire and pleasure are associated consistently with activity in mesolimbic brain regions, particularly the ventral striatum and ventral prefrontal cortex (Kuhn & Gallinat, ; Berridge & Kringelbach, ). Recent neuroimaging research demonstrates that the distinctiveness of different emotional categories is not as clear as once assumed (Hamann, ; Lindquist, Wager, Kober, Bliss-Moreau, & Barrett, ). However, we believe that on balance, evidence supports core neural constituents for specific emotions. Specifically, these emotions are distinguishable at the neurobiological systems level (which include both anatomical and neurochemical components) and are shared across individuals and species (Berridge & Kringelbach, ; Panksepp, Lane, Solms, & Smith, ). However, we also agree with perspectives put forth in traditional cognitive neuroscience that the interaction between older brain regions and higher neocortical struc-
we briefly discuss methodological approaches and current controversies, and close with important challenges for future research in this area.
Terms and Concepts Much of the terminology used herein refers to common concepts con cepts in psychology ps ychology and biology. Terms such as motivation and emotion refer to “states” that are generally understood but difficult to clearly define. ough some general aspects of these concepts are shared in the field, important differences in conceptualization can lead to misunderstandings among researchers. erefore, we offer some level of definitional detail and provide examples for key concepts in the following sections. Motivated Behavior Behavior
Motivated behaviors are focused and goal-directed. Motivated Importantly, however, not all “goal directed” behavior can be conceptualized as motivated behavior. Motivated Motiva ted behavior tends to be focused and highly
tures can generate complexities that are uniquely human (Panksepp et al., ). Additionally, we agree that emotional distinctions are not likely to emerge at the level of individual structures but rather in interactions of several regions across integrated circuits (Hamann, ). In this chapter, we consider the confluence of emotion and motivated behavior from from a neuroscientific and biological perspective. Our overriding framework is that, from the standpoint of behavioral expression, expression, behaviors that arise from emotion are motivated behavior (Beauchaine & Zisner, ). Behavior that emerges from emotional ex-
prioritized in terms of neuronal resources (Bradley et al., ). Such behaviors typically involve movements that are rapid, are direct, and contain an element of urgency (Beatty, Cranley, Carnaby, & Janelle, ). For example, walking across the street is a behavior that is clearly goal oriented, but it does not necessarily meet our definition. In contrast, if this behavior was done rapidly to escape the cold or to retrieve one’s crying toddler, it would be considered motivated. motivated. Given Given that motivated behavior contains an element of urgency, it may be performed in favor of other potential behavioral lures or influences (such as cross-traffic or encounters
perience directed, and focused by the emotionalis energized, brain systems engaged: for instance, withdrawal from threat is a behavior motivated m otivated by fear, the compulsive search for sex or drugs is behavior motivated by pleasure, and prolonged crying after losing a loved one is motivated by grief and sadness. us, motivated behavior and emotion are tightly coupled. However, they are not synonymous. ere are important ways in which emotion can be dissociated from motivated behavior, and some of these differences may be particularly important for psychological health and psychopathology. Wee begin by defining key terms and concepts, W then outline our theoretical perspective that emo-
with friends) in the the environment.
Emotion
Emotion is a difficult and often contentious concept to define clearly. Although terms such as fear and anger are commonly used in the scientific literature and offer convenient shorthand, interpretations of such terms vary. vary. Several theories have been proposed to explain the experience of emotion itself, referencing biological influences on physiological arousal (Scherer, ), cognitive labeling and contextual interpretation (Schacter & Singer, ; Reisenzein, ; Scherer, ; LeDoux, ; Barrett, ), and social learning processes (Fogel et al., ), among others (Barrett, ).
Whether such emotional concepts emerge from identical patterns of brain activation across individuals and species, require conscious experience, or are accompanied by universal expression are all important matters of debate (Ekman & Cordaro, ;
consequence of past experience. Memory is stored by novel patterns of brain activity and connections, which can elicit novel experiences that result from that past experience.
Damasio & Carvalho, ; LeDoux, ; Barrett, ; Panksepp et al., ). Working from a neurobiological perspective, we consider emotions to be coordinated patterns of activity in central and peripheral nervous systems, often accompanied by neuroendocrine activity (Panksepp & Biven, ). ese coordinated patterns of physiological activity are accompanied by experiential states that, at least among humans, are similar across individuals. Neural substrates of emotion within the central nervous system include both subcortical (deep and evolutionarily preserved across species) and neocortical (the outer layer of the brain that is the largest and most elaborated among primates and humans) regions. is activity is typically transitory—lasting seconds to minutes—and is often accompanied by some form of behavioral expression (Ekman & Davidson, ). ere are three important points to note about our conceptualization of emotion. First, a critical feature is coordinated activity activity of sub-
Attention
cortical and cortical structures. is generates internal states that are both multifaceted and organized. Second, compilations of brain activity associated with specific emotions are are recognizable recognizable patterns patterns but are not rigid. Just as the color red has many variations that remain “red,” fear has many variations from prototypical that still remain fear. ird, behavioral expression is a common but not necessary feature of emotion, which is particularly important for the subsequent discussion.
Social behavior refers to behavior (both affiliative and/or agonistic) that is targeted toward or coordinated with a conspecific.
Homeostasis Homeostasis is an important concept in physiology, and that has in or. biological studies ofone emotion andhistorical motivatedroots behavi behavior. Homeostasis is the active process of maintaining a steady state. For example, because maintaining constant levels of osmolarity is critical for life, many systems—both behavioral and physiological—function to maintain constant levels of fluidity and salinity within cells. ese processes may translate to human behavior: for example, the imbalance of fluids may lead to a sensation of thirst, which motivates the search for liquids to ingest.
Learning and Memory Learning refers to processes through which brain activation patterns and/or behavior change as a
Attention refers to active directing directing of sensory experience toward a specific domain, area, or sensory expectation.
Development Development refers to regulated maturational changes in the brain, body, and behavior that occur between birth and adulthood. Notably, although maturational changes in the brain follow a preprogrammed timeline or ontogeny, environmental experience and behavior play a critical role in guiding this process. One mechanistic example of this interactive process is neuronal pruning, where intrinsic signals guide exuberant initial synapse connection and circuit formation, which is then refined by removal of connections that are underutilized (Stiles, ).
Social Behavior
Teoretical Perspective Panksepp’s Framework A basic premise of the Pank Pankseppian seppian model is that emotions are “kinds” in the universe. Emotions arise from distinctive patterns of brain activity, conserved in their basic form across evolution, that function to generate highly motivated and evolutionarily adaptive behaviors. Because of their adaptive value, these organized patterns became embedded in thesystem genome and standard architectur architecture of the nervous (Panksepp & Biven, ).e Emotional kinds are brain systems organized organized primarily in subcortical structures in highly similar ways across both individuals and species. Panksepp identified seven basic emotional systems that that he believed were physiologically distinctive and form the basis of all emotional experience: SEEKING, which generates search and consummatory behavior; RAGE, which provokes provokes aggressive responses against conspecifics; FEAR, which is linked to withdrawal and avoidance behaviors; behavi ors; LUST, LUST, which generates mating matin g behavior; CARE, which leads to nurturant behavior; PANIC-GRIEF, which is associated with social separation and loss; and PLAY, which is related to , ,
affiliative exchange and promotes social cohesion and social learning (Panksepp & Biven, ). Each of these emerges from distinctive patterns of activity in subcortical brain systems and is present in similar form in most mammals. Some of these patterns are
tional experience, which can be construed as a meta-level of the primary process. is is frequently the level at which cognitive neuroscientists deal with emotion. e tertiary level involves engagement of representations of primary process experi-
even evident to varying degrees in birds, amphibians, and reptiles (Maclean, ; Panksepp, ). For example, the SEEKING system is a neural net work that ascends from the ventral tegmental area through the lateral hypothalamus and ventral striatum to the prefrontal cortex and consists primarily of dopaminergic fibers. e SEEKING system is associated with emotional states of wanting, excitement, and anticipation, and when engaged elicits motivated searches for elements needed for survival and procreation (Panksepp & Biven, ). An important important feature feature of Panksepp Panksepp’’s model is that emotions can be experienced at a variety of “levels.” e most basic and hard-wired is the primary process level. At this level, basic emotional systems can be most clearly differentiated and are most conserved across evolution. Panksepp considered primary process the raw feelings of emotions and argued that many traditional animal-based affective neuroscientists generally adopt this perspective in delineating
ences. According to Panksepp, secondary and tertiary process emotions would not exist without initial involvement of basic primary process states, but once they do appear they can function independently of primary process states. Panksepp and others argued that many psychiatric conditions are characterized by maladaptive emotional functioning at the secondary and tertiary process levels, in addition to the primary core levels (Panksepp, (Panksepp, ; LeDoux & Pine, ). is perspective is taken throughout this volume, and is a core aspect of the contemporary emotion dysregulation construct (Beauchaine, ). In addition to the three levels of emotional experience, another important aspect of the Pankseppian Pankseppian model is bidirectional communication. By bidirectional communication, Panksepp referred to brain responses that were both bottom-up (in which primary process emotions influence behavior and cognitive activity) and top-down (in which tertiary
the neural basis of emotional experience (Cahill & McGaugh, ; Moriceau, Raineki, Holman, Holman, & Sullivan, ; Orsini & Maren, ; Berridge & Kringelbach, ; Vanderschuren & Trezza, ; Panksepp et al., ). Although many brain-emotion researchers acknowledge a contribution of evolutionarily old subcortical systems to emotional experience, many argue that true affective experiences that humans consider “emotion” involve more complex cognitive processing. Such processing includes concepts such as consciousness, linguistic labeling, and sense of self (Rolls, ; LeDoux, ; Panksepp & Biven,
level emotional experience affects primary process activity). Panksepp argued that activity in both directions is stronger in humans than in other mammals because so much more of the human brain is composed of neocortex. is results in a greater dissociation between behavior and primary process emotions in humans. For example, neocortical activity can inhibit and potentiate primary process emotions, and also subvert or redirect (i.e., regulate) behavioral outcomes following primary process engagement. In the present context, this is an important observation because it indicates a greater degree of dissociation between emotional engagement and
; & Carvalho, ;). LeDoux, ; Barrett,Damasio ; Panksepp et al., Panksepp vehemently rejected this definition. However, he did argue that emotions consist of more than just core primary process states of intense activation. Panksepp argued that primary states generated at least two other levels of emotional experience. e secondary process emerges from the primary process as a function of learning. Consistent association of primary emotions with specific stimuli, classes of stimuli, or contexts can elicit partial activation of primary process emotions, mixtures of primary emotions, and even novel combinations of primary processes with novel brain activation patterns. Finally, Panksepp referred to a tertiary level of emo-
behavior humans than other animals. e ability to controlinbehavioral expression and regulate emotional experience has become an important adaptive skill in the modern human environment. Although Panksepp Panksepp’’s model has been very influential, it has not been accepted universally. ere are notable detractors and alternate interpretations of the neurobiology of emotion (Rolls, ; LeDoux, ; Damasio & Carvalho, ; Barrett, ; Panksepp et al., ). Indeed, aspects of the model, such as conscious experiential states of other species (Panksepp et al., ), are difficult to incorporate into standard scientific testing. However, because the model clearly incorporates mechanisms, brain function, and evolution into our discussion of
emotion more comprehensively than many other biological models, it is an important heuristic to which we appeal appeal for the remainder remainder of this chapter chapter..
Adaptive Nature Nature of Emotions and
(Rolls, ; Izquierdo & Murray, ; Saez, Rigotti, Ostojic, Fusi, & Salzman, ; Saez, Saez, Paton, Pa ton, Lau, & Salzman, ). Although directly relevant for homeostatic regulation, Panksepp argued that many functions associated with these neural
Motivated Behavior When viewed from the perspective of biological utility, there are several specific roles for emotion in promoting adaptive responses in fluctuating environments. Motivated behaviors are important intermediaries for many of these functions. We We highlight some of these specific functions in this section.
Homeostasis Because life is only possible within relatively strict parameters, active adaptations to fluctuations— both internal and external to the organism—are built into many aspects of physiology. Some of these operate at cellular and subcellular levels, but organism wide emotional behavior also plays an important role. For example, dehydration inevitably results in a motivated seeking response for water. One of the simplest demonstrations that emotional responses can index biological needs was demonstrated in a series of experiments on temperature hedonics. Cabanac () found that warm water baths of exactly the same temperature were rated as pleasant by those who were hypothermic but unpleasant by those who were hyperthermic, whereas the opposite was found for cool water baths. Tis simple experiment and many similar studies since demonstrated the functional feature of emotional systems in guiding behavior toward adaptive outcomes (Cabanac, ; Ramirez & Cabanac, ). Adaptive responding is also evident in experiments conducted by Curt Richter in the s in which he demonstrated that plasma reductions in sodium induced by adrenalectomy induced a highly specific motivated appetite for salt in rats (Richter, ). More recent examples include induction of panic attacks among individuals following following carbon dioxide buildup buildu p in the lungs (Pine et al., ), which presumably generates an energized search for consumable air. Panksepp viewed many of these homeostatic emotional responses as primary process behaviors, or ones that do not require learning and are not sculpted by environmental experiences.
structures relate to secondary process emotions. In contrast, primary process states are linked with environmental stimuli in an associative manner (Panksepp (Panksepp & Biven, ). Much of the interindividual variance in emotional tendencies is evidenced at the secondary process level: unique fear responses by different individuals following specific traumatic events are examples of secondary emotions being generated via associational processes. Tese secondary emotional associations therefore play an important role in many psychiatric conditions (Qin et al., ). Because environments vary widely between individuals, the ability to flexibly apply basic emotional activation patterns to unique environmental events is an important adaptive feature of motivational systems. Terefore, an important characteristic of emotional and motivational systems is that they capture attention and facilitate memory formation. Affective neuroscience studies indicate that emotion can capture attention by biasing limited neuronal processing of sensory signals toward those associated with the emotion with emotional al stimulus stimulus (Schet (Schettino tino,, Keil, Keil, Porcu, Porcu, & Muller, Muller, ; Hammerschmidt, Sennhenn-Reulen, Sennhenn-Reule n, & Schacht, ) via feedback mechanisms from the amygdala and possibly other regions to primary sensory areas (Pourtois, Schettino, & Vuilleumier, ). Tus, emotion may amplify relevant sensory responses and filter irrelevant sensory responses via associative regions including the amygdala and striatum. Emotion also acts as a “bottom up” means of facilitating acquisition of new behaviors and applying established response patterns to novel stimuli. Emotional Emotio nal facilitation of learning is a well-established phenomenon that applies to both appetitive and aversive events and affects conditional, instrumental, and social learning (Reisberg & Hertl, ). From Fro m the standpoint of biological adaptation, both the capturing of attention and learning are a means of employing motivated behavior in a maximally flexible manner across a number of different stimuli and contexts that may vary substantially across individuals.
Attention, Learning, and Memory Memory Several more recent studies of reward valuation and satiation in monkeys have focused on roles the amygdala and orbitofrontal cortex play in assigning value to stimuli with homeostatic importance
Social Behavior In social interactions, basic emotions direct motivated behavior in adaptive ways. For animals that live in social groups, emotion modulation plays an , ,
important role in guiding many aspects of behavior (Chang et al., ; Feldman, ). e PLAY, NURTURE, and LUST systems all involve positive, approach-related emotional motivation toward conspecifics. e RAGE and PANIC emotional
peaking shortly after puberty (Steinberg & Morris, ; S. J. Blakemore & Mills, ). From the standpoint of biological utility, developmental shifts in patterns of emotional responding may direct motivated behavior toward features or
systems engage motivated behavior in agonistic en counters and in response to separation distress. Although some specific behavioral patterns differ across mammalian species, there is considerable homology in neurochemical and neuroanatomical controls over affective responses related to these motivated behaviors. For example, there is a high degree of concordance in the controls of maternal behavior and sexual desire across mammals (Argiolas & Melis, ; Love, ; Lonstein, Levy, & Fleming, ). For humans and many nonhuman primates, the social world is of critical importance and also incredibly complex. Variables such as dominance, kinship, reciprocity, and alliance formation factor into every encounter (de Waal, ; Cheney & Seyfarth, ; Dunbar & Shultz, ). Emotionally mediated motivated behaviors play important roles in guiding many aspects of social behavior. Among humans, elaborate brain systems involved in perceptual and cognitive processing of social information are interconnected strongly with brain regions related to motivated behavior and emotion (Adolphs, ; Chang et al., ). is connectivity suggests that emotion and motivation permeate virtually all aspects of social behavior. behavior.
Development e role of motivation in development is often overlooked, but is a vitally important adaptive feature. e stimuli and contexts that induce motivated behavior vary in a systematic fashion across development (Nelson, Lau, & Jarcho, ; Nelson, Jarcho, & Guyer, ). is is perhaps most evident in changes that take place in social motivation during adolescence. However, However, regulated changes in affective sensitivity may apply in other domains and during other developmental periods as well (Forbes & Dahl, ; Spear, ; Spielberg, Olino, Forbes, & Dahl, ; Nelson et al., ). For example, caretaker separation induces a potent motivated response in early development, but this is greatly attenuated with age (Zhang et al., ). Similarly, Similarly, physical play with peers demonstrates a classic inverted-U response that peaks in late childhood and declines into adolescence and adulthood (Panksepp, Siviy, & Normansell, ; Vanderschuren & Trezza, ). Emotional responses to peer acceptance follow a similar trajectory,
stimulus categories that are most relevant for the specific phase of development the organism is in. Development of the nervous system includes mechanisms such as pruning, which is sensitive to environmental input. Because emotional responding directs attention and learning toward developmentally relevant features in the environment, through motivated behavior it may have potent effects on developmentall outcomes. developmenta
Maladaptivee Emotional Responses Maladaptiv Despite the fact that emotions evolved because of their capacity to generate biologically adaptive motivated behaviors, there are a number of ways in which motivated responses can be nonadaptive. Among humans, humans, one factor that that contributes contributes to maladaptive emotional experience and expression is the radical difference between environments that modern humans have constructed and our environments of evolutionary adaptation (Pinker, ). is is particularly evident in motivated behaviors associated with the SEEKING emotional system. Obtaining hydration, calories, nutrients, and social contact with group members both is essential for survival and depends on behavioral engagement. Appetitive behavioral responses responses geared geared toward toward seekseeking such essential features in the environment generate powerful reward responses upon consumption, which evolved in environments where sought-after stimuli such as high-calorie foods were scarce. However, such items are readily available now, making compulsory seeking and overconsumption much more problematic than in our evolutionary history. history. e most obvious example of this is overconsumption of food, but recent changes in the availability of constant social contact via social media provides another illustration of potentially problematic overconsumption (Primack et al., ; Shensa et al., ). Furthermore, the ability to cultivate and manufacture neurochemical agents (opiates, cocaine, methamphetamine) that potently activate the seeking/reward system independent of environmental experience has created a huge societal problem in which the functional utility of endogenous reward processes have, for some, been usurped (Panksepp, Herman, Conner, Bishop, & Scott, ).
A second factor that contributes to maladaptive emotional responding in humans is the extent of bidirectional communication between “top down” regulation and executive functions and “bottom up” primary emotional systems. Humans have developed a strong capacity to willfully regulate behav-
excessive alcohol consumption or behavioral shutdown that accompanies depression (Sharp, ). Although emotions evolved because of the adaptive value of motivated behavioral responses, resulting responses are not always optimal. is is particularly relevant for modern humans, in part
ioral expression and override emotional/motivational tendencies by engaging top-down inhibitory mechanisms (Buhle et al., ). is is often an adaptive response in certain environments/context environments/contextss and can serve as an important means of managing excessive emotional expression (Buhle et al., ). However, the ability to dissociate motivated behavioral expression from the experience of emotion can eventuate in dysfunctional brain response patterns in the long term. Chronic stress associated with the repeated overriding of natural motivated behavior tendencies predicts mood and anxiety disorders, and compensatory drug and alcohol use (Mah, Szabuniewicz, & Fiocco, ; Nusslock & Miller, ; Sharp, ). us, there are both benefits and potential drawbacks to using executive functions to override motivational tendencies. e maladaptive aspect of strong bilateral connections between primary emotions and executive functions is
due to the highly flexible nature of human behavior and executive functions, and in part the result of the relatively artificial environment humans have created for themselves.
also evidentsystems in reverse—hyperactivation basic emotional can generate long-termofalterations in executive functions such as attention and memory, thereby reinforcing dysregulated emotional expression (MacLeod et al., ; Rapee & Heimberg, ). Another way in which emotional/motiva emotional/motivational tional activation and behavioral responses can differ meaningfully is in the duration of experience. Whereas salient stimuli and events often occur transiently in the environment, emotional responses can linger from seconds to hours and generate even more long-lasting emotional states (Heller et al., , , ). e persistence of emotional responding in the absence of behavioral action can result in long-term detrimental functioning (Teicher, Samson, Anderson, & Ohashi, ), although emotion may be amenable to conscious regulatory intervention in the absence of behavioral output (Denny, Inhoff, Zerubavel, Davachi, & Ochsner, ). Finally, an important aspect of the relationship between emotion and behavior in humans is its equifinality: there is not always a unique behavioral expression for each experienced emotion. For instance, fear may generate drug seeking, compulsive hand-washing, or freezing. Anger may elicit agonistic physical encounters, but it may also trigger
pants engage emotionality in a task (usually a computer) that manipulates withon either standardized (e.g., expressive faces or monetary gains/losses) or customized probes (Salimpoor, Zald, Zatorre, Dagher, & McIntosh, ; Abrams et al., ). Although most neuroimaging approaches still rely on regression and parametric statistical comparisons, a number of novel approaches have emerged recently. ese include graph theory, causal modeling, independent component analysis, and machine learning approaches such as multivoxel pattern analysis (Bullmore & Sporns, ; Marinazzo, Liao, Chen, & Stramaglia, ; Saarimaki et al., ; Xie, Douglas, Doug las, Wu, Wu, Brody, Brody, & Anderson, Anderson , ). A key key problem problem with these these approaches approaches is that that they impose strong limits on the degree to which behavior can be expressed. Brain measures typically require data acquisition in a highly controlled environment in which participants are not allowed to move. However, some recent approaches have incorporated measures measures of subtle differences in behavioral responses expressed inside the scanning environment. ese differences may provide important insights into the relation between emotion and behavior. For example, researchers have begun to investigate brain differences in generating approach or withdrawal responses with joysticks (Radke et al.,
Current Methods and Findings ere are a number of ways to assess the effects of emotions on brain and behavior. In human-based studies standard approaches include functional neuroimaging, electroencephalography, and magnetoencephalography. More recent approaches include functional near-infrared spectroscopy (Hoshi, ), in which the blood oxygen level–dependent (BOLD) signal is assessed through light sensors placed on the skull, and transcranial electrical stimulation, in which current is delivered to the brain from scalp electrodes (Yavari, Jamil, Mosayebi Samani, Vidor, & Nitsche, ). e typical approach in all of these methods is to have partici-
, ,
, ), by directing attention toward or away from a threatening stimulus (Price et al., ; Ceravolo, Fruhholz, & Grandjean, ), and via grip strength elicited by aversive emotional stimuli (R. L. Blakemore, Rieger, & Vuilleumier, ). In some ways, animal studies are much more
using pattern analysis techniques have detected emotion-specific patterns in neuroimaging studies (Hamann, ; Kragel et al., ; Saarimaki et al., ; Nummenmaa & Saarimaki, —but see also Lindquist et al., ). Although emotion-specific differences in brain activation are detectable in
flexible for inducing emotional states and measuring brain and behavioral responses. In addition to standard pharmacological, electrophysiological, and targeted lesion methods, more recent advances include genetic knockouts (Crawley et al., ; Araragi & Lesch, ) and in vivo regulation of
human neuroimaging studies, these differences tend to be evident only at the network level and are not reflected in one-to-one mapping of brain regions with specific emotio emotions. ns. Although more specific associations between individual brain regions or neurochemical systems is more evident in animal-based
brain function with optogenetics (Deng, Xiao, & Wang, ). Human analogs of some of these ap Wang, proaches include assessment of behavior among individuals following naturally occurring brain lesions (Damasio & Carvalho, ), or in rare cases invasive measurement of brain function with intracranial recordings or deep brain stimulation, performed for clinical reasons (Pourtois, Spinelli, Seeck, & Vuilleumier, ). A full accounting of findings from research on the intersection between emotion and motivated behavior in the brain is beyond the scope of this chapter,, but general observations can be made based chapter
emotion research than in human research, it is clear that emotions are better characterized with networks than by activity within unique and specific neural structures (Floresco, ; Douglass et al., ).
ere continues to be a great deal of debate among investigators who study the neurobiology of affect regarding what exactly constitutes an emotion, to what extent different different emotions represent represent “kinds” “kinds” in nature, how similar emotional experience is across individuals, and how much homology exists in emotional experience across species. Panksepp’s af-
on findings.states First, in engageme engagement of emotional andexisting motivational humansnt and animals typically involves networks of brain regions including both subcortical and neocortical structures (Hamann, ; Kragel, Knodt, Hariri, & LaBar, ; Nummenmaa & Saarimaki, ). Sub(neo) cortical regions are more closely associated with what is typically considered to be the motivational (i.e., behavioral responsiveness) component of emotion, even though associated behavior is restricted in most human-based studies. e most prominent structures associated with motivation and emotion include the insular and anterior cingulate cortices (which are evolutionarily older cortical regions that lie underneath the neocortex), the amygdala, striatum, and midbrain regions such as the ventral tegmental area and periaqueductal gray. Aberrant activity in these regions is often associated with emotion/motivation-based psychopathologies— such as anxiety, depression, addiction, and borderline personality (Drevets, Savitz, & Trimble, ; Filbey et al., ; Lichenstein, Verstynen, & Forbes, ; Rigoli, Ewbank, Dalgleish, & Calder, ; Gola et al., ; Hein & Monk, ; Lago, Davis, Grillon, & Ernst, ; Sharp, ). Second, there is a tremendous amount of variability in patterns of brain network activity engaged
fective neuroscience model details unique signatures for seven basic emotional systems, each strongly tied to a range of motivated behaviors (Panksepp & Biven, ). In this model, different emotional systems are unique “kinds” in the brain: they are fundamentally shared across a number of species, typically contain specific behavioral outputs, and do not require high-level cognitive activity to be experienced. Although the details vary somewhat across investigators, this basic concept is largely shared by many researchers who adopt a traditional behavioral neuroscience and animal-based approach to emotion (Darwin, /; Maclean, ; Ekman & Davidson, ; Berridge & Kringelbach, ; Perusini & Fanselow, ; Panksepp et al., ). On the other end of the spectrum are researchers who argue against the notion that emotions are unique kinds in the brain, or that common experiences can be assumed across individuals and particularly across species (LeDoux, ; Barrett, ). Investigators Investigat ors who adhere to this perspective believe that emotion is largely a high-level cognitive interpretation that emerges from a variety of inputs, including but not limited to physiological response patterns. A variety of intermediary concepts lie along this continuum, including dimensional approaches in which emotion is construed as elaborations along
across studies. Howe However ver,, several recent meta-analyses
valence and arousal dimensions (Zachar & Ellis,
Ongoing Controversies
; Russell, ), and more truly hybrid approaches in which basic kinds exist but also interact with cognitive schemas and interpretatio interpretations ns (Izard, ; Damasio & Carvalho, ). Some of the controversy in this area relates to differences in semantics or fundamental inconsis-
advances are being made, and careful use of definition and terminology may further facilitate progress (Izard, ; Panksepp et al., ). Another issue that has plagued emotion research—particularly at the intersection of animal and human studies—is the role that conscious expe-
tencies in definitions and terminology (Izard, ). However, real conceptual differences do exist and have important effects on interpretation of the intersection of motivation and emotion. Although we certainly lean toward the Pankseppian Pankseppian model of basic kinds of emotion that are directly tied to
rience plays in emotional experience and expression (LeDoux, ; Panksepp et al., ). ere is no clear consensus on whether consciousness is necessary for emotion, whether conscious experiences are comparable enough across species to make generalizations, or even what the basic elements of con-
motivation and behavior, behavior, this is by no means a universally accepted approach (Barrett, ). Indeed, many of the initial neuroimaging studies that have attempted to differentiate emotional experiences based on functional brain activation have not been supportive of many of the specific models. Although functional neuroimaging has revealed emotional activation of both cortical and subcortical structures (Pauli et al., ; R. L. Blakemore et al., ), there tends to be a great deal of overlap between different emotions. Furthermor Furthermore, e, contrary to Panksepp’s model, imaging studies of emotions often do not demonstrate strong activation patterns
sciousness are. ese are issues at the forefront of neuroscience and philosophy—at present we have no clear empirical resolution.
with within brain stem brain regions reg ions (Haman (Ha mann, n, ; using ; Lindquist et al.,in). However, recent studies more sophisticated approaches to inducing and measuring emotion have reported unique patterns of activation by emotional category, largely contained within the cortex (Hamann, ; Saarimaki et al., ). An important caveat of all magnetic resonance imaging research is that, although it is the state of the art for noninvasively measuring neuronal activity,, it imposes several methodological constraints on ity the elicitation of emotion. Neural “events” must be generated while participants lie perfectly still inside a huge metal tube, and because the BOLD signal is relatively weak, experiences must be repeated many times over the course of a single experimental session. e emotional experiences induced under these circumstances are likely only a very weak resemblance to those generated by the naturalistic and highly salient situations typically associated with emotion. e experiences measured in the scanner may be closer to what Panksepp has referred to as secondary or tertiary emotional experiences derived from the more potent subcortically based primary emotions (Panksepp & Biven, ; ; Panksepp et al., ). Controversies such as this have plagued emotion research since its inception (Izard, ; Barrett, ) and continue to be a barrier for truly
all focusedthat on emerge a singlefrom stimulus, event, states or goal. Behaviors motivational are usually directed toward adaptive outcomes, although this is not always the case. Motivated behavior is usually a highly effective means of obtaining desired goals, but can be particularly difficult to manage or suppress when those goals are not adaptive for the individual. Indeed, emotion induction does not always generate behavior. e ability of an individual to manage behavioral expression in spite of emotional engagement, or the ability to manage the degree of emotional engagement elicited by a stimulus (emotion regulation), is an important skill that can also result in complex neuronal and psychological states that may be linked to psychopathology. ere are several important directions for future research to address. First, from a practical standpoint, it is important to gain a better understanding of how to manage motivated behavior that is maladaptive for the individual. Two classic examples of maladaptive motivated behavior are compulsive drug use that occurs in addiction and excessive avoidance behaviors that occur in anxiety. Probably the most effective way of altering such maladaptive motivation is through a combination of medication and/or top-down regulation to reduce the intensity of the motivational state and encourage behavior
integrative neuroscience research. However, some
that is inconsistent with the maladaptive motivation.
Teoretical Synthesis and Future Directions Our perspective is that emotion has strong and pervasive effects on behavioral expression, and that any behavior that results from emotional induction is motivated behavior by definition. When compared to behavior that does not result from emotion induction, motivated behavior is pervasive in that it involves a coordinated effort of cognitive, motor, neuroendocrine, and other physiological systems
, ,,
Modern therapeutic approaches for humans, such as cognitive-behavioral therapy, use these methods to promote effective management of maladaptive motivated behaviors. For these examples, therapeutic goals might include reducing drug seeking or stopping avoidance of feared stimuli. ese changes in behavior are extremely challenging to achieve, but
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repeated changes to the association between emotion and behavior can improve the likelihood of adaptive outcomes. Although effective methods have been developed on these fronts, continued efforts to refine and improve these approaches are needed. Second, at a more holistic level, motivational states can often be parsed into a pre-exposure period and a consummatory period. is is most commonly done for rewarding states (Berridge & Kringelbach, ; Luijten, Schellekens, Kuhn, Machielse, & Sescousse, ), but also may apply to aversive situations where exposure to a potentially aversive condition is anticipated before it is experienced (Grupe & Nitschke, ; Jarcho et al., ). In both appetitive and aversive conditions, the most powerful effects of motivation appear in the period preceding exposure. is is also where the maladaptive behavioral aspects of motivated behavior are mostcomponents prevalent. of ea motivated anticipatory and consummatory experience are fundamentally different from each other at a neurobiological level, but they are also completely interdependent. An important challenge for motivational research will be to gain a better understanding of how consummatory and anticipatory components of a motivated experience are regulated and how they vary across individuals. Finally, a fundamental question for motivation research is how novel motivational states become acquired. Although some stimuli (e.g., required nutrients, potential predators) are inherently motivational, for others the motivational states are acquired through learning (e.g., money) or development (e.g., sexual desire). Moreover, the motivational pull of stimuli that were once highly appealing can fade over time. Understanding how these motivational states change at the level of the brain and fluctuate with experience is an important feature of motivational learning that will be crucial to characterize further. further. References Abrams, D. A., Chen, T., T., Odriozola, P., P., Cheng, K. M., Baker Baker,, A. E., Padmanabhan Padmanabhan,, A., . . . Menon, V. (). Neural circuits underlying mother's voice perception predict social communication abilities in children. Proceedings of the National Academy of Sciences , ,, –.
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