Sample Chapter - Cognitive Neuroscience (4th ed.), Banich & Compton (2018)

Marie T. Banich and Rebecca J. Compton

COGNITIVE NEUROSCIENCE

Sample Chapter

Fourth Edition

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BRIEF TABLE OF

CONTENTS CONTENTS

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Preface Acknowledgments Dedication

xiv xvii xix

Part I Fundamentals Chapter 1 Introduction to the Nervous System Chapter 2 Historical Perspectives Chapter 3 Methods

1 2 41 68

Part II Neural Bases of Mental Functions Chapter 4 Motor Control Chapter 5 Sensation and Perception Chapter 6 Object Recognition Chapter 7 Spatial Cognition Chapter 8 Language Chapter 9 Memory and Learning Chapter 10 Attention Chapter 11 Executive Function and Higher-Order Thinking Chapter 12 Emotion Chapter 13 Social Cognition

99 100 136 167 198 223 256 296 332 367 394

Part III Broader Applications Chapter 14 Psychopathology Chapter 15 Brain Development and Plasticity Chapter 16 Generalized Cognitive Disorders Chapter 17 Cognitive Neuroscience and Society

421 422 455 491 522

Endpapers Glossary References Index

543 544 558 648

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Chapter 12 Emotion

By March 30, 1981, James Brady had been serving for several months as the press secretary for US president Ronald Reagan. Brady was known and liked by the White House press corps for his wit and energy. For example, during a lunch with reporters, he described a particular government bureaucrat as sleeping “in the closet hanging upside down with his wings over his eyes” (Bumiller, 1982). But March 30 was a terrible day for Jim Brady and all those who knew him. During a gunman’s attempt to assassinate President Reagan, Brady took a bullet in the head.

The injury was very severe – in fact, at one point that evening, CBS anchorman Dan Rather mistakenly reported that Brady had died (Bumiller, 1982). Eventually surgeons were able to reduce the swelling and bleeding in his brain so that Brady survived. However, because of the bullet’s trajectory, he suffered extensive brain damage to his right frontal lobe (Cytowic, 1981).

Many of the symptoms that Brady experienced are predictable from what we have already learned about the frontal lobes. Brady suffered paralysis of the left arm and leg, consistent with damage to the motor regions in the right frontal lobe. He also showed cognitive symptoms of frontal lobe damage, such as difficulties with initiating action and a tendency to perseverate in his thought.

But the gunshot wound did not just affect Brady’s cognitive functions – it affected his emotional regulation and emotional state as well. In an essay, his wife Sarah Brady wrote: [S]trong feelings of any kind could bring on what we called a “wail” – a very unnerving noise somewhere

Imagine a day without emotion. According to the psychologist and philosopher William James, living without emotion would require a person to “drag out an existence of merely cognitive or intellectual form.” James thought this rather undesirable, arguing that “[s]uch an existence, although it seems to have been the ideal of the ancient sages, is too apathetic to be keenly sought after by those born after the revival of the worship of sensibility” (James, 1884, p. 194). Though emotions can be destructive, they also bring vitality to our lives; they permeate nearly all aspects of our thoughts, decisions, and interactions with other people.

This chapter surveys different components of emotion and the brain systems that are important in implementing them.

First, though, we must be able to answer a basic question: What is an emotion? It has been said that everyone knows what an emotion is until they are asked to define it (LeDoux, 1996).

The American Heritage Dictionary of the American Language defines emotion as “1. Agitation of the passions or sensibilities often involving physiological changes. 2. Any strong feeling,

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Chapters begin with

between crying and laughing. As his brain healed, he was an opening case increasingly able to control it, and in later years, he would wail only during extremely emotionalhistory moments – to sad pique or happy – such as the singing of the national anthem.

But students’ interest in those early days, it happened all the time: He would preview start to say something, and suddenlyand his voice would just issues wail off. that are discovered

inasthe chapter At other times, though, Brady was later described speaking in a “slow, measured cadence” (De Witt, 1990) that lacked the emotional inflections of normal speech.

After his injury, Brady also tended to be a bit more brutally honest than people in the political sphere are generally inclined to be. For example, he made highly unflattering remarks about some of his former colleagues in the White House, sometimes making those around him a bit uncomfortable (Bumiller, 1982). Although these tendencies may simply reflect the change in outlook that accompanies a brush with death, they may also reflect a failure of his damaged frontal lobes to inhibit socially inappropriate behavior.

Despite his tragic circumstances, Brady continued to maintain his trademark wit; about John Hinckley, the man whose bullet hit him but missed President Reagan, Brady said, “I think that guy was an awfully bad shot.”

During the remainder of their lives, Jim and Sarah Brady dedicated themselves to advocating against gun violence and for the recognition of people with traumatic brain injuries. While James Brady can teach us about the emotional consequences of right frontal lobe damage, he also teaches us a lesson about emotional resilience in the face of tragedy. as of joy, sorrow, reverence, hate, or love, arising subjectively rather than through conscious mental effort.” At first glance, this description seems to capture our everyday sense of the concept of emotion. However, when we examine this definition more closely, we begin to appreciate the complexity of trying to understand emotion.

This definition assumes some things about emotion that still arouse heated debate in the scientific literature. For instance, what does it mean for a feeling to arise “subjectively rather than through conscious mental effort”? Indeed, much of the processing that is associated with emotion seems to occur outside conscious awareness. We may know how to describe a feeling we are experiencing, but we are often unaware of how that feeling was generated. Nevertheless, most of us would recognize that conscious mental effort can play a role in generating or maintaining emotions. Imagine, for example, the jealous lover who dwells on thoughts of his beloved in someone else’s arms, making himself more miserable in the process.

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Over 250 full-color images help students visualize and retain key information

Chapter 12 Emotion

Dilates pupil

Inhibits salivation

Oculomotor nerve (cranial nerve III)

Contracts pupil

Facial nerve (cranial nerve VII) Stimulates salivation

Glossopharyngeal nerve (cranial nerve IX)

Heart Accelerates heartbeat Relaxes airway

Slows heartbeat Constricts airway

Cranial

Lungs

Cervical

Liver Stimulates digestion Inhibits digestion Constricts blood vessels Inhibits digestion

Thoracic

Contracts bladder

Cranial Vagus nerve (cranial nerve X)

Cervical

Gallbladder Stomach Stimulates digestion

Thoracic

Spleen Stimulates digestion Pancreas Large intestine

Lumbar

Lumbar

Stimulates digestion Small intestine

Sacral

Sacral

Rectum Adrenal gland Coccygeal

Coccygeal Sympathetic chain ganglion

Kidney

Fight or Flight

Rest and Digest Stimulates bladder

Sympathetic Nervous System Stimulates orgasm

Parasympathetic Nervous System

Stimulates penile erection and labial engorgement

FIGURE 12.2 The autonomic nervous system.

Activation of the sympathetic branch of the autonomic nervous system is important in many bodily expressions of emotion, such as changes in heart rate, respiration, and sweat secretion. In contrast, the parasympathetic branch is activated under resting conditions.

NOTE: Pages throughout this sample chapter skip to give you insight to the key features in the book.

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Subcortical Contributions to Emotion Reward and Motivation

Key Just as identifying potential dangers is important for an organterms are ism’s survival, it is equally important for the brain to signal what situations and actions lead to rewards. Classic work on introduced the neural basis of reward and positive motivation dates back in boldface to the 1950s, when Olds and Milner (1954) carried out experand defined iments demonstrating that electrical stimulations to certain in a glossary parts of the brain were “rewarding” for a rat. But what do we mean when we say the rats found the stimulation “rewarding”? at the back The rats would press a lever hundreds and hundreds of times of the book to activate a current that stimulated specific brain regions.

Because the rats were willing to work so hard for this stimulation, the researchers inferred that it was rewarding. The dopaminerareas where stimulation is most rewarding are the dopaminer gic pathways stretching from the ventral tegmental area of the midbrain to a cluster of cells in the basal forebrain known as the nucleus accumbens (see Figure 12.7). This region is also referred to as the ventral striatum, because it is the ventral part of the basal ganglia (striatum).

It is tempting to refer to the reward pathway as the “pleasure center” of the brain, but caution is required. Just because an animal presses a lever repeatedly for stimulation, does that mean the animal gets pleasure from it? It is important to dissociate “wanting” something from actually “liking” it. This distinction is clear in research on addiction, which has demonstrated that addicts may go to great lengths to obtain a drug (i.e., wanting it), but no longer experience

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pleasure once they take it (i.e., liking it; Robinson et al., 2015). Some researchers have proposed that the dopaminergic path leading to the core of the nucleus accumbens is not responsible for pleasure itself, but for the “wanting” aspects of reward-related behavior – those aspects that propel an Medial forebrain bundle (MFB)

Substantia nigra Ventral tegmental area

Nucleus accumbens Lateral hypothalamus Amygdala Hippocampus

FIGURE 12.7 The location of the nucleus accumbens.

The nucleus accumbens receives ascending dopaminergic input from the ventral tegmental area.

Cortex

Core of the nucleus accumbens Corpus callosum Putamen

Nucleus accumbens

Shell of the nucleus accumbens

FIGURE 12.8 The nucleus accumbens core and shell regions in the rat brain.

The core of the nucleus accumbens is thought to play a role in wanting or desire incentive, whereas the shell is thought to play a role in consummatory pleasure. Courtesy of Harold Prüss.

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Chapter 12 Emotion

animal toward desired goals (Berridge and Robinson, 1998, 2003). In contrast, only a certain part of the nucleus accumbens – specifically, a layer of cells surrounding the accumbens and referred to as the nucleus accumbens shell – is thought to underlie the sensation of consummatory pleasure upon achieving a desired goal; that is, the “liking” (see Figure 12.8) (Berridge, 2003; Castro and Berridge, 2014; see also Baliki et al., 2013).

Many studies of the reward pathways have focused on nonhuman animals, but neuroimaging studies have also examined the conditions that activate the nucleus accumbens in humans.

The accumbens becomes activated in people when they receive a reward, especially when the reward is unexpected (Berns et al., 2001). In situations with predictable rewards, the accumbens is includes activated when the person anticipates the reward Each chapter before actually receiving it (Knutson et al., 2001). These results an “In Focus” box that suggest that the accumbens is initially sensitive to unexpected exploresrewards; in depth however, as a pattern of rewards emerges, the accumbegins to anticipate the reward (see also Fiorillo et al., a specificbens applied 2003; Hollerman and Schultz, 1998). issue in cognitive Evidence generally indicates that there is one basic brain neuroscience, system that processes rewards, regardless of the specific nature of those rewards (Berridge and Kringelbach, 2015). For examhelping students see ple, the nucleus accumbens is activated by a variety of stimthe implications of uli that could be considered rewarding, such as sweet juice, researchmoney, for real life faces (e.g., Aharon et al., 2001; Berns et and attractive al., 2001; Kampe et al., 2001; Knutson et al., 2000). This region

is also activated by rewarding items that are addictive. For example, smokers show greater accumbens responses to smoking-related imagery than do nonsmokers (David et al., 2005).

For obvious reasons, clinically oriented research on the ventral striatum has centered on its role in addiction, to which we will return in Chapter 14.

While the nucleus accumbens is most widely known for its role in reward processing, evidence suggests that, once again, a simplistic view is not consistent with all the evidence (Floresco, 2015). Instead, we should appreciate that the nucleus accumbens is well positioned to connect information about reward incentives to action systems. Intuitively, this makes sense: we don’t just feel craving for desirable chocolate, but this desire leads us to get up and root around in the cupboard until we find a tasty morsel of it. Through its anatomical connections with subcortical motor systems, specifically the basal ganglia, the nucleus accumbens aids us in reward-seeking, that is, in acquiring the desirable objects or experiences that meet our needs. Current evidence suggests that the nucleus accumbens is particularly important in biasing behavior toward seeking desired goals when obstacles arise or when there are many options from which to choose. Thus, the nucleus accumbens might not be necessary in enabling you to reach for a chocolate that is right in front of you, but it might prompt you to seek out the chocolate that is hidden on a top shelf, or to choose the chocolate from among the many other– but less delicious – options in your refrigerator.

In Focus: The Pleasure of Music What makes music so enjoyable to hear? No matter your specific musical preference or cultural upbringing, there is likely to be some kind of music that brings you pleasure. That pleasure may take the form of a transcendent state of bliss while listening to a classical piano concerto or a joyful urge to get up and move to your favorite dance tunes. Regardless, there is no doubt that music has a direct path to our emotions: it can bring us up out of our doldrums and connect us to something beyond ourselves. Studies of music cognition suggest that the temporal patterns of music – the rhythmic and melodic expectations that music establishes – can explain its pleasurable properties in part (Salimpoor et al., 2015).

Interestingly, even people who have great difficulty accurately perceiving musical pitch can still identify the emotions conveyed by music, perhaps because they can extract music’s temporal patterns (Gosselin et al., 2015).

Cognitive neuroscience studies further help us to understand how the interaction between auditory and emotion-related areas of the brain can facilitate the pleasure of music. First, recent research has detailed with high precision the ways in which music activates the temporal lobe regions involved in auditory processing. Participants in one fMRI study listened to a set of 165 different sounds, some of which were speech sounds, others which were either instrumental or vocal music (e.g., guitar, pop song), and others that were familiar sounds that were neither speech nor music (e.g., siren wailing, cell phone

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vibrating, water splashing; Norman-Haignere et al., 2015).

Researchers then applied a sophisticated computational analysis which, by examining profiles of activity across all voxels in the primary and secondary auditory cortex, identified clusters of voxels consistently activated by certain kinds of sounds. One cluster was specifically activated by speech and another was specifically activated by music.

To better understand the power of music to bring us pleasure, we need to consider how these auditory regions representing music interact with the brain’s reward pathways.

Music – especially music that we like– activates these reward pathways, specifically the nucleus accumbens (Koelsch, 2014).

Further linking musical pleasure with the reward system, one study found that dopamine release increased synchronously with participants’ peak emotional reactions to music (Salimpoor et al., 2011).

Putting these two lines of research together, a recent study examined connectivity between the auditory cortex and nucleus accumbens while people listened to music that differed in its level of pleasantness to the participant (Salimpoor et al., 2013).

After listening to unfamiliar musical segments, the participants indicated how much money they would pay for the music; in this way, researchers had an objective measure of the desirability of the music. Music deemed more desirable was associated with greater activity in the nucleus accumbens compared to music deemed less desirable. Moreover, when participants listened to

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Chapter summaries allow students to review the material learned or preview a loud clap of thunder while at the beach. You are likely to what to ofbe discussed engage in aissuite cognitive and behavioral changes associated with a feeling of fear. You may activate the fight-or-flight response, mentally note your own racing heart rate, remember the last time you got caught in a thunderstorm, alter your visual attention toward the skies and your auditory attention toward possibly new rumbles of thunder, attempt to regulate

Summary your feelings (“maybe that thunder is distant”), alter your facial expressions and tone of voice (assuming a worried expression and perhaps yelling, “Kids! Time to get out of the ocean!”), and make decisions about whether to pack up and go home or wait it out. Ultimately, any comprehensive understanding of emotional experience must take into account how all these aspects of emotion are integrated within the brain.

SUMMARY Subcortical Contributions to Emotion • The hypothalamus mediates some of the physiological phenomena associated with emotional states, such as changes in autonomic nervous system and endocrine function that are associated with fleeing or fighting.

• The amygdala is involved in learning the emotional significance of information and in producing a quick, instinctive, emotional response. The amygdala can also influence how attention is directed to emotionally significant events.

• The ventral striatum, or nucleus accumbens, is important in reward-seeking behavior. It is especially responsive to unpredicted rewards and becomes active when a person is anticipating a reward.

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Cortical Contributions to Emotion • The insula is involved in representing internal body states that are relevant to emotion. It is also important for the coding for unpleasant tastes, and also plays a role in the experience and perception of disgust.

• The anterior cingulate is involved in integrating information regarding emotion, cognition and action. It is also involved in evaluating the motivational outcome of actions.

• Orbitofrontal cortex is involved in assigning value to items and actions. It is also involved in evaluating reward and

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• •

punishment contingencies and in responding adaptively to changes in these relationships. Damage to the orbitofrontal cortex can lead to deficits in controlling (and exhibiting) behavior and emotion in a socially appropriate manner.

Control of emotions depends upon interactions among cortical and subcortical brain regions. Suppressing an emotion appears to involve top-down control over subcortical systems such as the amygdala and hypothalamus.

Temporoparietal regions of the right hemisphere are important for comprehending emotional information expressed in tone of voice or facial expression.

The right hemisphere plays a predominant role in the expression of emotion, both producing prosody that is related to emotional affect and in governing the expression of emotion on the face.

Positive affect is associated with more activity over the left than the right prefrontal cortex, whereas negative affect is associated with the reverse pattern (greater right prefrontal than left prefrontal activity).

States of high arousal appear to differentially involve the right hemisphere, particularly in posterior regions.

A particular emotional state may evoke activity across a widely distributed pattern of brain regions, because experiencing an emotion involves many components, including alterations in autonomic control, interpretation of bodily signals, actions, and alterations in attention and decision making.

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“An excellent introduction to the field of cognitive neuroscience – comprehensive, insightful and well written. The focus on translating neuroscience knowledge to clinical and societal issues is timely and sets the book apart. A great resource for anyone who wants to learn about the brain!”

Sabine Kastner, Professor of Psychology, Princeton University and the Princeton Neuroscience Institute

“Banich and Compton provide a broad, current, and accessible introduction to the mind and brain in Cognitive Neuroscience. A compelling introduction to the field.”

David Badre, Associate Professor, Brown University

“Cognitive Neuroscience details the impressive array of neuroimaging techniques available today alongside current findings. The engaging examples will readily captivate new scholars.​” Rachael D. Seidler, Professor, Applied Physiology & Kinesiology, University of Florida

Updated fully, this accessible yet thorough text highlights the most important theoretical, conceptual, and methodological issues in cognitive neuroscience. Written by two experienced teachers, the consistent narrative ensures that students link concepts across chapters, and the careful selection of topics enables them to grasp the big picture without getting distracted by details.

• Introduces students to the fascinating intersection of brain and cognition, and explores how the brain supports mental functions such as perception, memory, language, attention, emotion, and social cognition • Engages students and relates the material to real-world concerns through use of examples within the text, focus on clinical applications such as developmental disorders, brain injuries, and dementias, as well as opening case studies and “In Focus” boxes • Equips students with critical thinking skills that will enable them to critically evaluate new developments in the future • Summaries at end-of-chapter consolidate knowledge around key points New to this edition • All chapters have been extensively updated to include more current techniques, as well as a greater emphasis on how the brain works as a network • Includes a new chapter on neuroscience and society that considers how cognitive neuroscience issues relate to the law, education, and ethics, highlighting the clinical and real-world relevance of the field • The chapter on emotion and social processing has been split into two separate chapters, reflecting the growth in these areas of research • The illustration program has been updated to improve consistency and student understanding with more conceptual diagrams and figures to highlight major concepts Cover image: Sherbrooke Connectivity Imaging Lab (SCIL)

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