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Chapter 3

The Motivated and Emotional Brain

MOTIVATION, EMOTION, AND NEUROSCIENCE
Day-to-Day Events Activate Specific Brain Structures
Activated Brain Structures Generate Specific Motivations and Emotions

NEURAL BASIS OF MOTIVATION AND EMOTION

Cortical Brain
Subcortical Brain
Bidirectional Communication

INDIVIDUAL BRAIN STRUCTURES INVOLVED IN MOTIVATION AND EMOTION

Subcortical Brain Structures
Reticular Formation
Amygdala
Reward Center: Striatum, Nucleus Accumbens, and Ventral Tegmental Area
k Pleasure Cycle k
Addiction
Motivated Action: Basal Ganglia
Hypothalamus
Cortical Brain Structures
Insula
Prefrontal Cortex
Orbitofrontal Cortex
Ventromedial Prefrontal Cortex
Dorsolateral Prefrontal Cortex
Anterior Cingulate Cortex
HORMONES
Cortisol
Oxytocin
Testosterone
SUMMARY
READINGS FOR FURTHER STUDY

As you and a friend walk into the psychology building, you eye a poster recruiting volunteers for
a neuroscience experiment. Volunteers will receive $50 for their time, so you look at each other,
nod approvingly, and decide to give it a try. After all, neuroscience experiments tend to pay well.
Upon arrival, you see an experimental room that looks more like a hospital than a psychology
department. A huge machine—an fMRI scanner—occupies one room, a second fMRI scanner
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The Motivated and Emotional Brain 45

occupies an adjacent room, and lots of computers and screens occupy a third “master control” room
to monitor what is happening to the people who lie inside those two huge brain-scanning machines.
The experiment begins. You enter one fMRI machine and your friend enters the other.
Together, you will play a game and make decisions. The game will have two rounds. In round 1, you
will be the “proposer” and your friend the “responder.” To start the round, you are given $20 and
then asked how much you want to keep for yourself and how much you wish to give to your friend.
The rules are simple: If the responder (your friend) accepts your offer, you both keep the money
agreed upon; if the responder rejects your offer, neither keeps any of the money. This is a game that
pits self-interest (keep most of the money for yourself) versus social concern (be fair and share the
money).
In round 1, you propose $12 for yourself and offer $8 to your friend. After all, it is your $20 and
it is your decision what to do with the money. You send the message, “Keep $12, Give $8,” and wait
to see if your offer is accepted or rejected. A minute later, you see “Accept Offer.” You now have
$12, and your friend has $8.
In round 2, the tables are turned so that your friend is now the proposer and you are the responder.
You wait for an offer. The offer arrives: “Keep $19, Give $1.” This is not fair! This is not right! It is
greed and exploitation! Feeling a bit hot, you press “Refuse Offer.” You lose a free $1, but you also
stop your greedy so-called friend from taking advantage of you.
The experimenter invites you and your friend into the master control room so that she can show
your in-game brain activity. With a couple of clicks, several brain images appear on the computer
screen, as in Figure 3.1. Looking at the brain scans, she explains, “Okay, let me see if I can guess what

x=6 x = 54
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(a) (b)
Courtesy of Dr. Johnmarshall Reeve

x = 18 x = 33

(c) (d)
Figure 3.1 Brain scans of the “Proposer” in Round 1 and the “Responder” in Round 2
Note: Nucleus accumbens activations appear in the upper-left panel (a); dorsolateral prefrontal cortex activations appear in
the upper-right panel (b); amygdala activations appear in the lower-left panel (c); and insula activations appear in the
lower-right panel (d). All images are taken from fMRI imaging. In each picture, the front of the brain appears on the left

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46 Chapter 3 The Motivated and Emotional Brain

decisions you made.” She points to the brain scan on the upper-left side of Figure 3.1—you in the role
of the Proposer during round 1. She says, “I can tell from this activity in your nucleus accumbens
(a) that you found the $20 to be attractive, and I can tell from this activity in your dorsolateral
prefrontal cortex (upper-right panel) (b) that you exercised self-control over the temptation to keep
all the money for yourself. From what I see here on the screen, I’ll bet you shared the money.”
You nod.
Then she points to the brain scans on the lower-left side of Figure 3.1—you in the role of the
Responder during round 2. She says, “Oh my. I can tell from this activity in your amygdala (c) that
you felt rather upset—perhaps even angry—about the offer. And, I can tell from this insular activity
(lower-right panel) (d) that you experienced negative feelings. You were quite upset, no? You refused
the offer, didn’t you?” Again, you nod.

MOTIVATION, EMOTION, AND NEUROSCIENCE

Neuroscience is the scientific study of the nervous system—and the human brain in particular. This
chapter, however, is not about the nuts-and-bolts biology of the nervous system (e.g., neurons,
synapses). Instead, the chapter looks at the human brain in action—how its neural structures and
pathways are associated with psychological processes (cognitive neuroscience; Gazzaniga, Ivry, &
Mangun, 2008), motivational processes (motivational neuroscience; Reeve & Lee, 2012), and emo-
tional processes (affective neuroscience; Davidson & Sutton, 1995).
The human brain is the marvel of the universe. There really is nothing else like it. It con-
tains approximately 3 pounds of gray matter (neurons and synapses) and white matter (connecting
fibers and axons) that are home to 100 billion neurons that support 100 trillion neuron-to-neuron
connections.
Why is the brain important? Most people say that the brain is important because it carries out
k cognitive and intellectual functions, including thinking, learning, remembering, decision making, k
and problem solving (Behrens, Fox, Laird, & Smith, 2013). Others, including physicians and those
who work in special education, say the brain is important to understand clinical conditions, such as
autism, dyslexia, and stuttering. Still others study the brain as a sensory-perceptual organ, because
much of what the brain does is process and make sense of the environment (e.g., the huge occipital
lobe processes vision). These are important brain processes, but the brain does more. The brain is also
the center of motivation and emotion. It generates cravings, needs, desires, preferences, pleasure and
pain, liking and wanting, and emotions and feelings. This chapter is specifically about the motivated
and emotional brain.

Day-to-Day Events Activate Specific Brain Structures

When someone flashes you a warm smile, a small brain structure in the back, right side of the
brain—the posterior superior temporal sulcus (pSTS) processes what you are seeing and rather
instinctively (automatically) knows that the person is happy and friendly (Srinivasan, Golomb, &
Martinez, 2016). Similarly, when someone flashes you an angry facial expression, the amygdala
processes this and rather instinctively knows that threat and danger are coming. In the chapter’s
opening vignette, the opportunity to gain money activated the nucleus accumbens, and it was the
injustice of being taken advantage of that activated the amygdala and insula. What these examples
illustrate is that environmental events in the physical and social world—events that range from preda-
tors, bullies, and loud noises to a pleasant smell, a humorous video clip, and finding a $20 bill that
we thought we had lost—produce brain activations.
In the laboratory, researchers understand brain functioning by stimulating specific brain struc-
tures to see what happens. They ask questions such as, “If I apply a mild electrical stimulation to
this particular brain area, what will happen?” In doing so, they can isolate the specific functions

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Neural Basis of Motivation and Emotion 47

associated with the various brain structures (e.g., “The amygdala does this, the nucleus accumbens
does that … ”). During surgery, for instance, the surgeon might apply a probe to send a mild electric
current to a specific brain structure. Since the brain has no pain receptors, such brain stimulation is
actually painless. When the surgeon stimulates one area of the brain, the person may move his or her
index finger. When the surgeon repositions the probe to touch another area, a particular sensation or
a childhood memory may come to mind.
Unless you are a surgeon, direct stimulation to the brain is a rather impractical idea. The current
gold standard to look inside the brain is functional magnetic resonance imaging (fMRI). The fMRI
detects changes in brain activity as active brain areas are fueled by glucose and oxygen. Glucose
and oxygen are both carried in the blood, and when a brain area becomes active, then blood—and
hence the glucose and oxygen within it—flows toward it. So, while a person is lying inside the fMRI
scanner, the neuroscientist can expose the person to some environmental stimuli (e.g., an emotional
facial expression) and follow the oxygen to observe the brain at work. There are actually several ways
to observe brain activity in real time, including measuring electrical brain activity directly with an
electroencephalogram (EEG) or measuring metabolic effects of changes in glucose absorption with
a positron emissions tomography scanner (PET scan). Whether the researcher uses fMRI, EEG, or
PET, this ability to observe and measure the live brain in action has led to an explosion of knowledge
about the neural bases of motivation and emotion.

Activated Brain Structures Generate Specific Motivations and Emotions

Half of the equation in a neuroscientific understanding of motivation and emotion is to understand
how internal (thought) and external (environmental) events activate specific brain structures (e.g.,
What activates the insula?). The other half of the equation is to know how specific brain structures,
once activated, energize, direct, and sustain motivational and emotional states (e.g., What does the
k activated insula do?). Different brain structures, when stimulated, give rise to specific motivational k
and emotional states that help us cope with and adjust to what is happening. Stimulating one part of
the hypothalamus, for instance, increases hunger, while stimulating a different part of the hypotha-
lamus increases satiety (feeling full). Increased neural activity in the anterior insula is necessary for
empathy (Gu et al., 2012), while increased neural activity in the dorsolateral prefrontal cortex is nec-
essary to exert self-control over temptation (Hare, Camerer, & Rangel, 2009). It is findings like these
that led neuroscientists to map out an understanding of which specific brain structures and pathways
are the neural basis of specific motivations and emotions.

NEURAL BASIS OF MOTIVATION AND EMOTION

The brain is astonishingly complex (Bassett & Gazzaniga, 2011). But considered at its most gen-
eral level, the brain features an outer cortical region and an inner subcortical region, as illustrated
in Figure 3.2. The cortical region is sometimes referred to as the cerebral cortex, and it generally
includes the bulges and grooves of the frontal, parietal, temporal, and occipital lobes of the brain.
The subcortical region is sometimes referred to as the limbic system, and it generally includes the
small nuclei that make up the anatomic core or center of the brain.

Cortical Brain
The outer cortical brain—the cerebral cortex (or “cerebrum”)—is the bulging, wrinkled surface that
most people think of when they think of the brain. It functions at a conscious, deliberate, intentional,
and purposive level (Szpunar, Watson, & McDermott, 2007). As such, the cortical brain is associated
with cognitively rich motivations such as goals, plans, strategies, values, and beliefs about the self.
The cortical brain is active as you set a goal to make an A on your psychology test, formulate a plan

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48 Chapter 3 The Motivated and Emotional Brain

Cortical
Region

Subcortical
Region

Figure 3.2 Subcortical and Cortical Regions of the Brain

of how you will attain that goal, adopt a strategic decision to read the book chapter on Thursday
evening, and exercise the self-control needed to resist the temptation to binge watch Netflix.

Subcortical Brain
The subcortical brain is associated with basic urges and impulses and with emotion-rich motivations
such as hunger, thirst, anger, fear, anxiety, pleasure, desire, reward, and wanting. For instance, when
you are at the airport and pass by one of those shops selling hot, fresh cinnamon rolls, the alluring
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aroma stimulates your subcortical brain to rather automatically generate an urge to approach the
source of the felt pleasure. These urges and emotions occur regardless of whether you want them
to—hence, these motivations are largely unconscious, automatic, urgent, and impulsive.

Bidirectional Communication
The brain features many individual structures, but it is important to note that these individual struc-
tures are linked together by a network of neural pathways (i.e., white matter). That is, almost all
individual brain structures project out nerve fibers that act as information superhighways to commu-
nicate reciprocally with other brain structures. These communication pathways allow the cognitive,
motivational, and emotional states that arise in one area of the brain to inform, contribute to, and
change the cognitive, motivational, and emotional states that arise in another part of the brain. So,
activity in one brain region causes upstream activity in another brain region (“I’m so mad at her,
I could yell.”), while activity in the second brain region causes downstream activity to modify the
original brain activity (“She probably didn’t do that on purpose. It is no big deal. I’m not so angry
anymore.”).
The overall picture of brain function is therefore not one in which individual brain structures are
associated with individual functions (e.g., the amygdala does this, the nucleus accumbens does that)
but, rather, one of interconnectivity and the brain working in a highly integrated way (O’Doherty,
2004).
Motivational and emotional states can be generated in a “bottom up” fashion as individual
brain structures respond to environmental stimuli (sweet taste, having your hand restrained against
your will). These motivations are typically generated by the subcortical brain. Similarly, the corti-
cal brain continually exerts “top down” processes (cognitive appraisal of an event, as in “Oh, this

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Individual Brain Structures Involved in Motivation and Emotion 49

is good news” or “Oh, this is bad news.”) that generate and modify motivational and emotional
states. So, in interacting with the world, the subcortical brain generates motivational and emo-
tional states and, in thinking about and making sense of the world and our place in it, the cortical
brain generates and modifies motivational and emotional states so that subcortical and cortical brain
structures sometimes work together (e.g., “Yes, this is good, let’s do it.”) but other times work in con-
flict (e.g., we hear the dentist drill and begin to feel some pain coming on, but we nevertheless stay
in the chair telling ourself that our visit is actually a good, not a bad, thing that is happening to us).
Here is an example of bidirectional communication. Imagine seeing a smiling face on a person
who is walking toward you and that person is a member of a distrusted out-group (Paulus & Wentura,
2017). The subcortical brain will rather automatically generate approach motivation in response to
the smiling face, while the cortical brain will rather reflectively generate avoidance motivation in
response to knowledge about what membership in the distrusted out-group means. To experience a
coherent motivational–emotional state and to forge a plan of action, there needs to be a great deal
of bidirectional communication between the cortical and subcortical brain. So, we scan the person’s
face, appearance, and pace for more information, and we scan our memories of what has happened
in the past in similar situations, as we try to integrate the conflicting information into a viable plan
of action.
Subcortical brain regions concern basic motivational processes (e.g., “Ice cream—I want it!”),
while cortical brain regions concern matters such as self-control, resisting temptation, decision
making, assessing risk, and self-regulation. The bidirectional forces between basic motivations
and cognitive control over these basic motivations and emotions has been termed the dual-process
model. Dual-process models are especially informative in understanding motivation, additions,
and risk-taking during childhood and adolescence. During childhood, subcortical brain processes
(reward-driven affective impulses) tend to dominate the cortical brain and its reflective cognitive
capacities, because childhood is an age in which the cortical brain structures are still developing and
k maturing (Best, Miller, & Jones, 2009; Cragg & Nation, 2008). For instance, adolescents take more k
risks than do adults, at least in the use of alcohol, tobacco, legal and illegal drugs, dangerous driving,
unprotected sex, and criminal behavior (Arnett, 1991). The basic neurological problem underlying
adolescent risk-taking is that mature subcortical brain structures are hot and actively involved in
decision making, whereas immature cortical brain structures are cold and less actively involved
in decision making (Galvan, 2010; Galvan et al., 2006; Somerville, Hare, & Casey, 2010). This
research suggests an overall picture in which the affective subcortical brain and the cognitive
cortical brain are two interacting systems (i.e., dual processes) that are sometimes in competition
and conflict with each other (Gladwin, Figner, Crone, & Wiers, 2011). With greater development
and maturation of the cortical brain, children, adolescents, and young adults become increasingly
able to control strong motivations and emotional processes (e.g., urges, impulses, addictions) and
to delay immediate gratification for the benefit of long-term goals.

INDIVIDUAL BRAIN STRUCTURES INVOLVED IN MOTIVATION AND EMOTION

Sixteen brain structures closely associated with motivational and emotional states are mapped
anatomically in Figure 3.3. The figure is the sort of image produced by an MRI or fMRI machine.
One structure lies within the brain stem (the final upper portion of the spinal cord)—the reticular
formation. Nine structures reside in the subcortical brain—amygdala, ventral striatum, nucleus
accumbens, ventral tegmental area, hypothalamus, caudate nucleus, putamen, substantia nigra,
and globus pallidus. Six structures reside in the cortical brain—insular cortex, prefrontal cortex,
orbitofrontal cortex, ventromedial prefrontal cortex, dorsolateral prefrontal cortex, and anterior
cingulate cortex. To complement the anatomical focus of Figure 3.3, Table 3.1 identifies the key
motivational and emotional functions associated with each of these 16 brain structures.

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50 Chapter 3 The Motivated and Emotional Brain

(a) (b) Dorsolateral Prefrontal Cortex

Ventral
Anterior Striatum & Globus
Cingulate Nucleus Pallidus Caudate Nucleus
Cortex Accumbens
& Putamen

Hypothalamus
Prefrontal
Cortex Substantia
Nigra

Insular Cortex

Caudate Nucleus

Courtesy of Dr. Johnmarshall Reeve

Putamen

Orbitofrontal Insular Cortex

Cortex
Ventral Globus Pallidus
Ventromedial Tegmental
Prefrontal Cortex Area
Amygdala

Reticular Formation
(c)

k Figure 3.3 Anatomical Locations of the 16 Key Motivation- and Emotion-Related Subcortical and Cortical k
Brain Structures

Subcortical Brain Structures

Reticular Formation
The reticular formation plays a key role in arousal, alertness, and the process of awakening the brain
to process incoming sensory information. The reticular formation is a cluster of neurons within the
brain stem about the size of your little finger. It is located at the apex of the spinal cord where the
spinal cord ends and the subcortical brain begins. It consists of two parts: the ascending reticular
activating system and the descending reticular formation. The reticular activating system projects
its nerves upward to alert and arouse the brain, whereas the descending reticular formation pro-
jects its nerves downward to regulate the body (e.g., muscle tonus). It is the reticular activating
system that wakes, alerts, and arouses the brain to ready it to process the incoming information.
Once aroused, the alerted brain processes the incoming information (e.g., makes a decision about
what to do) and, a second later, responds with appropriate action and coping.

Amygdala
The amygdala (meaning “almond-shaped”) is a collection of interconnected nuclei associated
with emotion and motivation (Baxter & Murray, 2002; McDonald, 1998). One primary function
of the amygdala is to automatically and instantaneously detect, learn about, and respond to
emotionally significant and aversive events, although each of its different nuclei serves a different
function. Stimulation of one part of the amygdala generates emotional anger, while stimulation of
another part generates emotional fear and defensive behavior (Blandler, 1988). Also, impairment

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Individual Brain Structures Involved in Motivation and Emotion 51

Table 3.1 Motivational and Emotional Function of the 16 Specific Brain Structures Featured in Figure 3.3

Brain Structure Motivational or Emotional Function

Subcortical Brain
Reticular formation Arousal, alertness, wakefulness.
Amygdala Detects, learns about, and responds to the stimulus properties of environmental
objects, including both threat-eliciting and reward-eliciting associations.
Basal gangliaa Motivational modulation of movement and action.
Striatum and nucleus The brain’s reward center. Responds to signals of reward (dopamine release)
accumbens to produce pleasure, wanting, liking, and approach.
Ventral tegmental area Starting point in the brain’s dopamine-based reward center.
Manufactures and releases dopamine.
Hypothalamus Responsive to natural rewards in the regulation of eating, drinking, and mating.
Regulates both the endocrine system and the autonomic nervous system.
Cortical Brain
Insular cortex (insula) Monitors bodily states to produce gut-felt feelings. Processes feelings
associated with empathy, intrinsic motivation, risk, uncertainty, pain, and
personal agency.
Prefrontal cortex Making plans, setting goals, formulating intentions. Right hemispheric activity
is associated with negative affect and “no-go” avoidance motivation, while
left hemispheric activity is associated with positive affect and “go” approach
motivation.
Orbitofrontal cortex Stores and processes reward-related value of environmental objects and events
to formulate preferences and make choices between options.
Ventromedial prefrontal Evaluates the unlearned emotional value of basic sensory rewards and internal
cortex bodily states. Responsible for emotional control.
k Dorsolateral prefrontal Evaluates the learned emotional value of environmental events and possible k
cortex courses of action. Responsible for control over urges and risks during the
pursuit of long-term goals.
Anterior cingulate cortex Monitors motivational conflicts. Resolves conflicts by recruiting other cortical
brain structures for executive or cognitive control over basic urges and
emotions.
a The basal ganglia include the caudate nucleus, putamen, substantial nigra, and globus pallidus.

of these same amygdala nuclei will produce striking changes, including an overall tameness,
affective neutrality, a lack of emotional responsiveness, a preference for social isolation over social
affiliation, and a willingness to approach previously frightening stimuli (Aggleton, 1992; Kling &
Brothers, 1992; Rolls, 1999). These studies make it clear that one key function of the amygdala is to
generate stimulus-emotion associations related to self-preservation, such as fear, anger, and anxiety
(Hamann, Ely, Hoffman, & Kilts, 2002). If there is an aversive, emotionally charged stimulus in the
environment, the amygdala will detect and respond to it.
The amygdala detects environmental threat and generates threat-elicited defensive responses
(Cardinal, Parkinson, Hall, & Everitt, 2002; Gallagher & Chiba, 1996). Fear is the conscious real-
ization of threat-elicited bodily reactions such as heart rate acceleration, muscular tension, behavioral
freezing, and “fear face” facial expressions (LeDoux, 2013). As the person encounters and detects
a threatening object, amygdala stimulation occurs and activates neighboring brain structures (e.g.,
hypothalamus, ventral tegmental area) that release neurotransmitters (dopamine, serotonin, nora-
drenaline, acetylcholine) to instigate and regulate a coordinated defensive response, including rapid
breathing (Harper, Frysinger, Trelease, & Marks, 1984), heart rate acceleration (Kapp et al., 1982),
and high blood pressure (Mogenson & Calaresu, 1973), as well as hormonal discharge and emo-
tional facial expressions (Davis, Hitchcock, & Rosen, 1987). What the amygdala does is (1) detect

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52 Chapter 3 The Motivated and Emotional Brain

the aversive characteristics of environmental objects and (b) relay this emotion-laden information to
neighboring cortical and subcortical brain regions.
As one point of illustration, a rat with a lesioned amygdala will crawl all over a sleeping cat
and even nibble playfully on the cat’s ear (Blanchard & Blanchard, 1972). What is missing from
the fearless rat is its capacity to generate the hard-wired amygdala-coordinated defensive response.
Without an amygdala, the rat lacks the means to respond emotionally to the cat, and it also lacks
the capacity to learn to fear the cat when the cat wakes up and poses a threat. When humans have
their amygdala removed (to control epileptic seizures, for instance) they become calm, docile, and
emotionally indifferent, even in the face of provocation (Aggleton, 1992; Ramamurthi, 1988).
The amygdala processes the aversive and threatening characteristics of all environmental
objects, but it has a special skill in detecting the aversive and threatening characteristics within
facial displays of emotion (Adolphs, Tranel, Damasio, & Damasio, 1994; LeDoux, Romanski, &
Xagoraris, 1989; Rolls, 1999). If another person lowers her brow and presses her lips tightly
together (as when angry), the amygdala automatically, effortlessly, and reliably picks up on this
threat-eliciting information.
A second primary function of the amygdala is to detect, respond to, and learn about rewarding
and beneficial properties of various environmental objects and events (Baxter & Murray, 2002). What
amygdala nuclei detect, learning about, and respond to is the presence versus absence of reward, the
value or quality of the available reward, the predictability of the reward, and the costs associated with
trying to obtain the potential reward (Berridge & Kringelbach, 2008; Whalen, 1999, 2007). These
studies make it clear that a second key function of the amygdala is to generate stimulus-emotion asso-
ciations related to reward. If there is an attractive, emotionally charged stimulus in the environment,
the amygdala will detect it, evaluate it, and respond to it.
An example to communicate the overall threat- and reward-detecting function of the amyg-
dala can be illustrated by a person gambling. In gambling tasks, people need to be able to assess
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risk, weigh the cost and benefits of choices, and cope with changing outcomes and odds on those
outcomes. With each win, the amygdala generates positive emotion and relays that positive emotion-
ality to the ventromedial prefrontal cortex so that the person can make an informed judgment about
what to do on the next trial, considering the risks, choices, and changing probabilities. Similarly, with
each loss, the amygdala generates and relays negative emotion to the ventromedial prefrontal cor-
tex so that the person can again make an informed judgment about future decisions and behaviors.
In this case, the amygdala generates the emotional joy of winning and reward and the emotional
pain of losing and danger, while the ventromedial prefrontal cortex uses this emotional information
to make a judgment about future predicted winning and losing. However, people with damage to
their amygdala behave in a bizarre fashion during such a gambling task, because they have no fear
of risk and therefore act recklessly (Bechara, Damasio, Tranel, & Damasio, 1996). It is good to know
when to hold them and when to fold them, and it is the emotionally rich amygdala—not the rational
cortical brain—that knows best when to stay and when to quit.
The amygdala has an interesting anatomical relationship with other brain areas. The amygdala
sends projections to almost every part of the brain, although only a small number of projections
return information to the amygdala. This imbalance helps explain why emotion, especially nega-
tive emotion, tends to overpower cognition more than cognition tends to regulate emotion. Hence,
a lot of fear and anger messages get blurted out. This is because amygdala nuclei are mostly evo-
lutionarily old structures that produce primitive emotionality. However, the lateral amygdala nuclei
have undergone relatively recent development to forge reciprocal projections and pathways with the
cortical brain, especially with the frontal lobes (Cardinal et al., 2002). These evolutionary new path-
ways allow for some degree of conscious regulation of these biologically basic primitive emotions.
The overall picture is one of bidirectional communication, though the amygdala’s upstream messages
of fear, danger, and reward are stronger and more numerous than are the cortical brain’s downstream
messages of self-control and reflective thought (e.g., a second opinion on what is happening).

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Individual Brain Structures Involved in Motivation and Emotion 53

Reward Center: Striatum, Nucleus Accumbens, and Ventral Tegmental Area

The brain’s reward center consists of several subcortical brain structures that communicate with each
other through the dopamine network. Dopamine is a neurotransmitter that is crucial for motivation
and movement. As shown in Figure 3.4, the dopamine network begins with the ventral tegmental
area, continues with the striatum and nucleus accumbens, and ends in the basal ganglia to translate
reward-based motivation into action.
The striatum consists of the nucleus accumbens, caudate nucleus, and putamen (Liljeholm &
O’Doherty, 2012). The activation of the ventral (lower part) striatum is practically synonymous with
the experience of reward, or what neuroscientists term the “hedonic evaluation of stimuli.” Through
their activation, we learn what to like, what to prefer, and what to want (Smith, Tindell, Aldridge, &
Berridge, 2009).
Reward is fundamental to motivation. It is fundamental to survival, to learning, to well-being,
and to the generation of goal-directed effort (Schultz, 2000). When a person encounters an envi-
ronmental object (e.g., orange juice), its stimulus characteristics are processed in the amygdala and
ventral striatum (sweet taste, cool temperature), and the experience of rewarding and pleasurable
feelings occurs in the nucleus accumbens (e.g., “I like it.”) (Pecina & Berridge, 2005). The nucleus
accumbens is active during the experience of a pleasant taste, a pleasant image, social acceptance,
and several addictive drugs (Berridge & Robinson, 1998; Sabatinelli et al., 2007; Wise, 2002).
What specifically stimulates the nucleus accumbens to become active—what constitutes
“reward-related information”—is the release of the neurotransmitter dopamine (Berridge &
Kringelbach, 2008). Once activated by the release of dopamine, the ventral striatum translates the
experience of reward into motivational force, approach behavior, and the exertion of physical effort
(Pessiglione et al., 2007).
From a neuroscience perspective, reward is dopamine release. As people go about their day,
k some level of dopamine is always present in the brain. But as people encounter a variety of events, k
those that signal reward—a pleasant image (looking at a beautiful face), a pleasant taste (sipping
sweet juice)—trigger dopamine release (Sabatinelli et al., 2007; Wise, 2002). These triggering events
can be natural or learned rewards.
The ventral tegmental area is the manufacturing site for brain dopamine, so it is the starting
point in the brain’s dopamine-based reward center. The ventral tegmental area projects fibers into
the nucleus accumbens that receive dopamine-release information to create the biology (and the
experience) of reward. As shown in Figure 3.4, the ventral tegmental area-to-nucleus accumbens
pathway extends further upstream into the cortical brain. It is in the prefrontal cortex that the person
has a conscious experience of pleasure, and it is the orbitofrontal cortex that stores the learned reward
value of environmental objects so that the person will know (will remember) that a particular object
has produced rewarding consequences in the past.
Activation of the ventral tegmental area-to-nucleus accumbens dopamine pathway is what
allows people to learn the reward value of environmental objects and events. People learn the reward
value of any such object or event through stimulus appraisal (amygdala, ventral striatum) and then
through extent of dopamine release (ventral tegmental area, nucleus accumbens) (Hampton &
O’Doherty, 2007; Hayden, Nair, McCoy, & Platt, 2008; McClure, York, & Montague, 2004;
O’Doherty, 2004). The ventral tegmental area also relays reward-related excitatory signals to the
basal ganglia, which, in turn, send excitatory signals to initiate motivated action (i.e., approach the
reward-related event). Overall, the more dopamine that is released, the greater will be the learning,
positive emotion, approach motivation, and effort exertion.
When day-to-day events unfold in ways that are better than expected, the ventral tegmental
area releases increased dopamine, and the dopamine surge signals that the event is producing more
reward than it was anticipated to deliver. In contrast, when events unfold in ways that are worse than
expected, a decreased dopamine release serves as information that a particular course of action is
producing less reward than it was anticipated to deliver (Montague, Dayan, & Sejnowski, 1996).

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54 Chapter 3 The Motivated and Emotional Brain

Prefrontal
Cortex

NA
Orbitofrontal
Cortex
VTA

© Oguz Aral/Shutterstock

Figure 3.4 The Dopamine-Based Reward Circuit

The brain’s reward system. The dopamine-based reward circuit begins in the ventral tegmental area (VTA)
where dopamine is manufactured and then released to the nucleus accumbens (NA). From the NA, the reward
k center extends into the prefrontal cortex, which is involved in the subjective experience of pleasure, and into k
the orbitofrontal cortex, which stores the object’s learned reward value.

When you anticipate good news or when you anticipate an exciting event, dopamine release
occurs. It is not the good news or the event itself that causes the ventral tegmental area to release
dopamine but is, instead, the anticipation of rewarding news and the anticipation of a rewarding event.
That is, the ventral tegmental area releases dopamine and the nucleus accumbens is activated when
we first learn that we are about to receive some money (reward anticipation), not when we actually
receive the money (reward receipt). Dopamine release is therefore greatest when rewarding events
occur in ways that are unpredicted (“Wow, I’m surprised how nice that flower smells.”) or under-
predicted (“Wow, that flower smells much nicer than I thought it would.”) (Mirenowicz & Schultz,
1994). For this reason, we typically experience more pleasure in thinking about eating chocolate chip
cookies and about engaging in sex than we do when actually munching on the cookies or engaging
in sex. Of course, if things go better than expected during the eating or mating, then the dopamine
release continues and so does its corresponding positive feeling and approach motivation.
How the dopamine system contributes into motivated action and how the various brain struc-
tures that are central to the dopamine system interact bi-directionally with subcortical and cortical
brain structures can be seen in Figure 3.5. The right side of the figure provides a summary of the
reward-based network of subcortical structures, while the left side of the figure foreshadows the
discussion to come on cortical brain structures and how they contribute into the dopamine-centric
reward system.

Pleasure Cycle
The extent of pleasure felt during motivational states rises and falls over time (Berridge
& Kringelbach, 2008, 2015). As shown by the solid line in Figure 3.6 (based on Kringelbach
& Berridge, 2017), behavior begins with initiation, as we explore for food at the grocery store or look

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Subcortical Brain Structures

Amygdala
Responsive to reward and threat characteristics of objects.

Hypothalamus
Responsive to natural rewards (e.g., food, water, and mating). Dopamine System
Motivated Action
Ventral tegmental area
Insula
Manufactures and releases dopamine to send Basal Ganglia
Represents bodily-based feelings.
forward the reward-related information Preparation for action.
about the environmental object. The caudate nucleus, putamen,
substantia nigra, and globus
Cortical Brain Structures pallidus project into the pre-
supplemental, supplemental,
Prefrontal Cortex
Ventral Striatum and Nucleus and motor areas to
Positive emotion and approach; negative emotion and
Accumbens motivationally and emotionally
avoidance.
Responsive to dopamine signals of reward infuse action.
Orbitofrontal Cortex in terms of pleasure and liking.
Store, process, and input the reward-related value of objects.

k Ventromedial Prefrontal Cortex k

Evaluate and input the unlearned (natural) reward value
of objects.
Dorsolateral Prefrontal Cortex
Evaluate and input the learned (acquired) reward value of
objects.

Anterior Cingulate Cortex

Monitor conflict. Exert cognitive control over decision making.

Figure 3.5 Neural Core of Dopamine-Centric Reward-Based Motivated Action

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56 Chapter 3 The Motivated and Emotional Brain

Typical Pleasure Cycle

Muted Pleasure Cycle from Addiction

Wanting Liking Learning

Strong Dominates Dominates Occurs

Amount of
Pleasure

Weak

Searching Consuming Satiation

Time
Figure 3.6 The Pleasure Cycle. The solid line represents the typical wanting–liking–learning cycle, while
the dashed line represents the muted pleasure cycle dominated only by wanting (without liking and learning)

for someone interesting to talk to at a party. Pleasure is low at this point in the pleasure cycle, while
wanting is high. If an attractive stimulus is found, it is consumed. At this point in the pleasure cycle,
pleasure and “liking” are high. Finally, consumption of an attractive object produces reward. At this
point in the pleasure cycle, liking declines while a strong learning effect is produced (i.e., one
learns where the good food and the interesting people are). So, the pleasure cycle is characterized,
in order, by wanting, then liking, then learning.
k Wanting is a motivational state that comes from an actual need for something (e.g., when dehy- k
drated, people want water). Liking, however, is a motivational state that comes from experiencing
pleasure (Berridge & Robinson, 1995). Wanting and liking typically go hand-in-hand (i.e., we want
and like the same things), but the two motivational experiences can diverge (Berridge & Robinson,
1998; Dickinson & Balleine, 2002), and this is what typically occurs during addiction. In Figure 3.6,
the solid line illustrates the naturally occurring wanting–liking–learning sequence, while the dashed
line represents the abnormal loss of liking that typically occurs during an addiction state.

Addiction
At first, smoking a cigarette, eating a sugary donut, or ingesting cocaine produces liking. But, because
pleasure is linked to dopamine-release and because dopamine-release occurs mostly with unexpected
reward, the liking-associated pleasure associated with the cigarette, donut, or cocaine inevitably
fades (because the reward becomes expected, rather than unexpected). At this point, wanting can
occur without liking—the person can physiological need (i.e., want) the addictive drug, though it no
longer produces pleasure (i.e., liking).
For those dopamine-generating rewards that have addictive properties (e.g., cocaine, heroin,
amphetamine, alcohol, and nicotine), the dashed line in Figure 3.6 better represents the muted plea-
sure cycle. Wanting without liking occurs when the nucleus accumbens becomes hypersensitive to
dopamine stimulation (Di Chiara, 1998). When addictive substances are used repeatedly, their liking
wears off (reward shifts from unexpected to expected) while the wanting only grows (because the
nucleus accumbens becomes hypersensitive to dopamine stimulation). This state of wanting without
liking can last for years (Hyman & Malenka, 2001; Robinson & Kolb, 1997).
To help smokers quit the habit, some currently marketed pharmaceuticals block the initial
dopamine-related pleasure typically generated by nicotine (e.g., the drug Chantix). These prescrip-
tion drugs prevent much of the nucleus accumbens-based “liking” (by taking the pleasure out of

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Individual Brain Structures Involved in Motivation and Emotion 57

nicotine intake). The reason why drugs such as Chantix can be moderately successful is because
individuals suffering from addiction often turn to the substance to temporarily boost their positive
mood state, especially when under the influence of stress or other such negative affect (Baker et al.,
2004; Carmody, Vieten, & Astin, 2007; Valentino, Lucki, & Van Bockstaele, 2010).
This pattern of suffering negative feelings (e.g., stress, loneliness) but looking for a temporary,
escape-based boost in positive mood can be seen in drug and alcohol abuse (Baker et al., 2004;
Carmody et al., 2007; Holahan et al., 2001; Kassel, 2010), but it can also be seen in gambling,
risk-taking, and excessive Internet usage (Griffiths, 2000; Leith & Baumeister, 1996; Valentino et al.,
2010; Young, 2004).
One illustration of “excessive Internet usage” can be seen in Facebook usage (Sheldon,
Abad, & Hinsch, 2011). “Facebooking” has become near epidemic in college populations (Pempek,
Yermolayeva, & Calvert, 2009). College students (and others) log on to this popular social network
site because it offers an easy opportunity to experience positive feelings and a sense of connection.
But a simulated social experience is not as deeply rewarding as is an actual face-to-face social
interaction. What the research shows with frequent Facebook usage (logging on more than twice
a day) is that Facebooking does somewhat alleviate negative feelings, but it also paradoxically
reduces longer-term positive affect and well-being (because the user experiences fewer face-to-face
interactions; Sheldon et al., 2011). Overall, frequent Facebook users may become somewhat
“hooked” (addiction is probably too strong a word) on an easily accessible, mild, short-term boost
in positive affect that can distract them from negative feelings, and this is especially true for a new
user (because the positive affect boosts is new and relatively more unexpected). But Facebooking is
problematic as a coping mechanism to negative feelings when the lonely person goes to Facebook to
substitute virtual interactions for face-to-face interactions and relationships (Sheldon et al., 2011).
The key condition that generates a deep sense of relatedness satisfaction, social connection, and
interpersonal intimacy is how responsive one’s interaction partner is to your needs and concerns
k (Reis, 2014). For the person looking for deeply satisfying, happy, and rewarding relationships, the k
place to find these experiences is in the real world and in a relationship partner who cares deeply
about, is highly responsive to, and is unconditionally supportive of your welfare.

Motivated Action: Basal Ganglia

The essence of motivation and emotion is energized and persistent goal-directed behavior. Motivated
people move and take action. Movement and action flow out of neural activity in the motor cortex.
The motor cortex sends “go” signals to the body’s muscles to produce movement. Before such move-
ment occurs, however, the presupplemental and supplemental motor areas first plan, excite, inhibit,
and enact these motor commands. The presupplemental and supplemental motor areas, which are
located at the very top of the head (where you might pat a young child on the head), largely send the
motor instruction to the premotor and motor cortex and are therefore more related to movement and
action than they are to motivation and emotion per se.
Basal ganglia (basal meaning at the base of the cortex, ganglia meaning a group of nerve cells)
are a cluster of many different small nuclei in the subcortical brain that collectively provide move-
ment and action with a motivational and an emotional punch. The substantial nigra and globus
pallidus motivationally and emotionally prepare action; they make a planned action more or less
potent (more or less energized or invigorated). They are active, for instance, for the game show con-
testant as she eagerly presses the answer buzzer and for just about anyone in pursuit of rewards and
gains. The caudate nucleus and putamen give rise to movement intentions and coordinated (rather
than conflicted) action. All basal ganglia—substantial nigra, globus pallidus, caudate nucleus, and
putamen—are closely connected to and receive information from the cortical areas of the brain
(to receive action plans) and to the motor, premotor, supplemental motor area, and presupplemental
motor areas (to execute and carry out those action plans). The collective role of the basal ganglia is
to energize (or inhibit) those action plans (Pessiglione et al., 2007).

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Hypothalamus
The hypothalamus is a small subcortical brain structure that comprises less than 1 percent of the total
volume of the brain. Despite its small size, it is a motivational giant.
The hypothalamus exists as a collection of 20 neighboring and interconnected nuclei that serve
separate and discrete functions. Through the stimulation of its 20 separate nuclei, the hypothalamus
regulates a range of important biological functions, including eating, drinking, and mating (via the
motivations for hunger, satiety, thirst, and sex that are the subject of Chapter 4). The hypothalamus
is responsive to natural rewards (e.g., food, water, and sexual partners).
The hypothalamus also regulates both the endocrine system and autonomic nervous system.
By regulating these two systems, the hypothalamus is able to regulate the body’s internal environ-
ment (e.g., heart rate, hormone secretion) in order to adapt optimally to the environment (e.g., cope
with a stressor).

BOX 3 How and Why Antidepressant Drugs Alleviate Depression

Question: Why is this information important? to the source from where it came. The antidepressant acts
Answer: To understand how antidepressant drugs to restore serotonin levels and usage to normal.
alleviate depression. Depression is also associated with a diminished
capacity to experience pleasure—an inability to experience
Each of us wages a lifelong struggle against depres- pleasure and positive feelings for life events. Low dopamine
sion, and about 1 in 10 of us suffers from clinical levels can leave the person vulnerable to apathy, boredom,
depression. Disappointment, loss, failure, hassle, financial poor concentration, and with little initiative to embrace the
k woe, interpersonal neglect, and social rejection represent day. In contrast, dopamine release generates good feelings, k
the all-too-common flow of human experience. When such positive affect, and essentially leaves the person primed to a
events affect us in the moment, we feel sad or distressed. positive mood (Ashby, Isen, & Turken, 1999). Researchers
When such events affect us chronically, however, we feel have not been able to produce dopamine-based antide-
depressed. Aversive, stressful life events affect our bodily pressive drugs because dopamine decays too rapidly for
biochemistry and, when they deplete our biochemical pharmaceutical usage.
resources, they can leave us vulnerable to depression. Overall, depression has two faces: serotonin defi-
Depression is a complex psychological disorder ciency, which leaves the person less able to cope with
associated with the stress of coping and a diminished life’s stress, and dopamine deficiency, which leaves the
capacity to experience pleasure, or what clinical psychol- person less ready to anticipate and experience pleasure.
ogists refer to as “anhedonia.” Relative to the stress of Drugs targeting serotonin and dopamine both unfortunately
coping, exposure to uncontrollable stress makes demands produce troubling side effects. Antidepressant drugs not
on the subcortical brain that can gradually deplete brain only supply serotonin, but they also “hijack” dopamine
serotonin (a neurotransmitter). We need serotonin for pathways to some degree and inadvertently blunt feelings of
the motivation to cope, so serotonin deficiency leaves us
love, romance, and attachment to others (Zhou et al., 2005).
vulnerable to depression (Kramer, 1993; Weiss & Simson,
Few people taking antidepressants, for instance, have the
1985). The popular antidepressant drugs (e.g., Prozac,
experience of falling in love. Some addiction-countering
Zoloft, Paxil, Cymbalta, Lexapro) are SSRIs, or selective
drugs (e.g., smoking cessation) blunt dopamine release in
serotonin-reuptake inhibitors. These antidepressants work
on the premise that depression is caused by low turnover in general (and not just while taking the addictive substance)
the serotonin pathways. During stressful life events, sero- and therefore leave the depressed person a bit more vulner-
tonin is released into the synapse, but it also quickly returns able to suicide. Knowing this, pharmacological researchers
(experiences reuptake) to the sending neuron. To reverse are now working on a third alternative: namely, drugs that
depression, the antidepressant drug prevents serotonin produce neurogenesis, or the growth of new nerve cells.
update and hence makes serotonin more readily available, Sprouting new nerve growth that can generate positive affect
as it stays in the synapse area rather than returning (uptake) is one key biological event that keeps depression at bay.

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Individual Brain Structures Involved in Motivation and Emotion 59

The hypothalamus regulates the endocrine system by exerting control over the pituitary
gland—the so-called master gland of the endocrine (or hormonal) system (Agnati, Bjelke, & Fuxe,
1992; Pert, 1986). Anatomically, the hypothalamus is immediately north of the pituitary gland,
and it regulates the pituitary gland by secreting hormones into the tiny capillaries that connect the
hypothalamus to the pituitary gland.
The pituitary gland, in turn, regulates the endocrine system. For instance, to increase arousal, the
hypothalamus stimulates the pituitary gland to send hormones through the bloodstream to stimulate
the adrenal glands to release its own hormones (epinephrine, norepinephrine) into the blood stream
that trigger various bodily organs to initiate the well-known “fight-or-flight” response. A later section
in this chapter (“Hormones”) will explain how the hypothalamus regulates both the pituitary gland
and the stress hormone of cortisol (see Box 3).
The hypothalamus also controls the autonomic nervous system (ANS), which includes all
neural innervations into body organs that are under involuntary control (e.g., heart, lungs, liver).
It is divided into the excitatory sympathetic system that accelerates bodily functions and alerts the
body (as through an increased heart rate) and the inhibitory parasympathetic system that facilitates
rest, recovery, and digestion. Therefore, the autonomic nervous system begins at the hypothalamus
(the hypothalamus is the autonomic nervous system’s head ganglion, or starting point) and extends
its nerves throughout the body to generate both arousal (sympathetic activation) and recovery
(parasympathetic activation).

Cortical Brain Structures

Insula

k The insula is a rather large and highly interconnected structure that lies deep within the brain. k
Anatomically, it is the fold that lies between the posterior part of the frontal lobe and the anterior
part of the temporal lobe and also just above the subcortical brain (see Figure 3.3, image c).
The insular cortex (or insula) consists of two roughly equal halves—an anterior and a posterior
part. The posterior insula receives, monitors, and becomes aware of changes in bodily states such as
changes in heart rate, fatigue, temperature, touch, muscle tone, arousal, and cravings (Craig, 2003,
2009). It is more aligned with the subcortical brain, as many of these bodily based states lie beneath
conscious awareness. The anterior insula monitors, evaluates, and consciously represents (becomes
aware of) the subjective feelings that arise from these changes in bodily states. Hence, the anterior
insula monitors and becomes aware of “gut” (bodily based) feelings. It is more aligned with the
cortical brain.
The insula processes interoceptive information about the state of one’s body (visceral, homeo-
static), and it therefore allows the person to mentally construct a consciously aware representation
of how he or she feels (Craig, 2009; Wicker et al., 2003). When people have “a feeling about that
thing” (e.g., this person is untrustworthy, my homework is boring, class is enjoyable), it is activity
in the anterior insular cortex that gives rise to this feeling. So, the first key function of the insula is
to receive, process, and allow the person to become aware of “raw feelings,” gut-felt feelings, and
intuitive hunches (e.g., “women’s intuition”).
Pain is one bodily feeling experience that the insular monitors, but insular activity seems to be
involved in practically all subjective feelings (Craig, 2009), including not only negative feelings but
positive feelings as well (Gu et al., 2013). The right anterior insula processes “energy-consuming”
negative emotions (e.g., pain, disgust, anger, fear), while the left anterior insula processes
“energy-nourishing” positive emotions (e.g., pleasure, happiness, satisfaction) (Craig, 2011). It is
also in the anterior insula that people consolidate their internal bodily feeling state information with
external social-contextual information about the task they are involved in at the moment and the

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60 Chapter 3 The Motivated and Emotional Brain

social context in which they are in to form a basis of the conscious experience of emotion or affect
during that task (Craig, 2002, 2009).
The anterior insula is the key brain structure involved in intrinsic motivation (Lee & Reeve,
2017). The opposite of intrinsic motivation, which is extrinsic motivation, is the seeking and con-
suming of environmental rewards (e.g., food, money, social approval), and extrinsic motivation is
well explained by the striatum-based reward center depicted in Figure 3.4. In contrast, intrinsic
motivation arises from “intrinsic rewards,” such as subjective feelings of interest and enjoyment
(Lee, 2017). These intrinsic rewards are the “spontaneous satisfactions” one feels while engaged in
a task (e.g., satisfaction from a job well done), and it is this sense of task-generated satisfaction that
allows the activity to be experienced as interesting, enjoyable, and fun. These intrinsic satisfactions
are generated by the anterior insula (see the left side of Figure 3.7). As shown on the right side of
Figure 3.7, when people engage in intrinsically motivating activities, the more “intrinsic satisfac-
tion” they feel, the more they will show high levels of anterior insula activity (Lee & Reeve, 2013).
What this means is that when a person engages in a task “for fun” or “because it is interesting,” the
task creates anterior insular stimulation that generates a sense of satisfaction.
We will discuss intrinsic motivation in detail in Chapters 5 and 6, but the important point to
make here is that when one’s engagement in a task allows the person to feel curious, competent,
autonomous, or relatedness, it is activity in the anterior insula that is generating these feelings of
intrinsic reward.
The anterior insula is not only involved in the processing of one’s own feelings, but it is also
involved in the processing of the feelings of others. The anterior insula is the key brain structure
involved in empathy, which is the ability to perceive and share another person’s emotional state
(Gu et al., 2012). If you see another person in pain, you too will feel that pain, at least to the extent
that anterior insula activity occurs while you are observing the other’s pain. If anterior insula activity
does not occur while you observe another in pain, then you will likely not experience empathy
k for that other person. That is, anterior insula activity is necessary for an experience of empathy k
(Gu et al., 2012). This finding has led to some interesting speculation that a key neurological deficit of
people who lack a capacity for empathy is an insensitive or a damaged anterior insula—for instance,
people with autism, conduct disorder, or borderline personality disorder.
The insula also processes and learns about risk and uncertainty (Huettel, Stowe, Gordon, Warner,
& Plat, 2006; Kuhnen & Knutson, 2005). This is important because the role of the insula seems to
be to integrate current feelings, a risk prediction forecast (that always has a degree of uncertainty
associated with it) that arises from considering the consequences of one’s actions, and contextual

0.3
Extent of Anterior Insula Activity

0.2
x = –32
Courtesy of Dr. Johnmarshall

0.1

0
1 3 5 7
–0.1
Reeve

r = .79
–0.2
0 t = 10 Extent of Felt Satisfaction

Figure 3.7 Anterior Insular Activity Correlates with Extent of Task-Generated Satisfaction

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Individual Brain Structures Involved in Motivation and Emotion 61

information to produce a global feeling state that guides decision making (Singer, Critchley, &
Preuschoff, 2009). Much of that global feeling state exists as anxiety (Paulus & Stein, 2006). The
decision to trust another person, for instance, is one such instance of subjective feelings, risk, uncer-
tainty, considering the consequences of one’s actions, and decision making. When a generally coop-
erative person begins to act in a way that seems exploitive or untrustworthy (recall the chapter’s
opening vignette), insular activity occurs. The person picks up on social-contextual cues to experi-
ence a gut-felt feeling that “something is not right.” This feeling then enters into the decision-making
process whether to continue to trust that person. This same anterior insular activity occurs during
financial decision making as well, as in judging the risk and uncertainty of an investment (Kuhnen
& Knutson, 2005).
The insula is also responsible for a feeling of “self” and a sense of having a boundary that allows
for an intuitive distinction between me (self) and others (not self). Insular activity is also responsible
for an intuitive distinction between “action caused by me” and “action not caused by me” (Farrer
et al., 2003; Farrer & Frith, 2002). In an experiment, a participant will be asked to perform a simple
action (e.g., move a joystick), while the experimenter manipulates what happens when the person
performs that simple action. In one case, the participant’s action will cause a consequence (moving
the joystick makes an image appear on screen), but in another case the consequence will occur at
random (when the image appears is controlled by the experimenter, not the participant). In the first
case, the person will show insular activation and will experience “self-as-cause.” In the second case,
the person will not show insular activation and will experience “other-as-cause.” Hence, anterior
insular activity during action increases an experience of personal agency (Lee & Reeve, 2013).
Personal agency (I can change my environment in an intentional way) is fundamentally important
to motivation, because people are volitionally motivated to act when they feel “self-as-cause,”
because they believe their actions produce desired effects, but people are not so volitionally
motivated to act when they feel “other-as-cause” (Bandura, 2006). With “self-as-cause” personal
k k
agency, people willingly act on their environmental surroundings to change things for the better, but
with an “other-as-cause” lack of personal agency, people withhold effort because it seems rather
pointless.

Prefrontal Cortex
The prefrontal lobes of the cerebral cortex lie immediately behind the forehead. One lobe is on the
right side of the brain (right prefrontal cortex), while the other is on the left side of the brain (left
prefrontal cortex). Together, these two cortical lobes underlie many important motivations, including
affect, goals, and personal strivings.
The starting point for many (but not all) negative emotions is the amygdala, while the starting
point for many (but not all) positive emotions is the dopamine-network (ventral tegmental area and
nucleus accumbens). The phrase “but not all” is important, because emotions also arise from cortical
processes such as thoughts, appraisals, and goals.
The right lobe of the prefrontal cortex generates negative emotion and “no-go” avoidance moti-
vation, while the left lobe of the prefrontal cortex generates positive emotion and “go” approach
motivation (Davidson, 2004). If you watch a film clip showing puppies and babies (to induce pos-
itive emotion), you will show greater left than right prefrontal cortex activity; but if you watch a
film clip showing heartache and suffering (to induce negative emotion), you will show greater right
than left right prefrontal cortex activity (Fischer et al., 2002). This leads to an important conclu-
sion about the role of the prefrontal cortex in emotion: Left activations signal positive emotion and
approach motivation; right activations signal negative emotion and avoidance emotion (Davidson,
2012). More specifically, left prefrontal cortical activity is associated with parasympathetic nervous
system activity, calmness, positive emotionality, approach motivation, and group-oriented desires
such as affiliation, while right prefrontal cortical activity is associated with sympathetic nervous

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62 Chapter 3 The Motivated and Emotional Brain

system activity, arousal and danger, negative emotionality, and individual-oriented desires such as
personal protection.
Asymmetry in the prefrontal lobes leads to two interesting findings. The first is that people gen-
erally have greater activity in one lobe than in the other. This is assessed with EEG recordings when
the person is in a resting state. People with relatively greater left asymmetry (the left prefrontal cor-
tex is chronically more active than is the right prefrontal cortex) tend to engage in approach-oriented
behaviors such as reward-seeking, impulsivity, and aggression-dominance; people with relatively
greater right asymmetry tend to engage in avoidance-oriented experiences such as fear, depression,
and internalization symptoms (Davidson, 2004; Harmon-Jones, 2003). The second is that prefrontal
cortex asymmetry can not only be trait-like (as above), but it can also be manipulated and hence
state-like. People who squeeze a rubber ball with their right hand for a couple of minutes, for instance,
will show greater left prefrontal lobe activity and report a more positive mood, while people who
squeeze a rubber ball with their left hand will show greater right prefrontal lobe activity and report
a more negative mood (Harmon-Jones, 2006; Harmon-Jones, Gable, & Peterson, 2010).
The prefrontal cortex houses a person’s conscious goals (Miller & Cohen, 2001). Thoughts,
intentions, goals, and strivings that stimulate the left prefrontal cortex generate positive and
approach-oriented feelings, whereas thoughts, intentions, goals, and strivings that stimulate the
right prefrontal cortex generate negative and avoidance-oriented feelings (Gable, Reis, & Elliot,
2000). The associated positive versus negative emotion then colors which goals and strivings
the person does and does not pursue. The right prefrontal cortex is a cortical bathtub of negative
emotion in which thoughts, goals, intentions, memories, and personal strivings bathe as they
ready themselves for action—which usually takes the form of anxiety, caution, pessimism, and,
hence, avoidance, while the left prefrontal cortex is a cortical bathtub of positive emotion in which
thoughts, goals, intentions, memories, and personal strivings bathe as they ready themselves for
action—which usually takes the form of hope, eagerness, optimism, and, hence, approach.
k Because different people show different levels of sensitivities to process information in their k
right versus left prefrontal cortex, it means that biologically basic personality differences exist
between people that open them up to optimism, positive emotionality, and approach motivation
in their day-to-day thinking and planning (people with more sensitive left prefrontal lobes), while
other people are open or vulnerable to pessimism, negative emotionality, and avoidance motivation
in their day-to-day thinking and planning (people with more sensitive right prefrontal lobes; Gable
et al., 2000). An active and sensitive left prefrontal cortex provides the person with a behavioral
activation system (BAS), which is similar to extraversion, while an active and sensitive right
prefrontal cortex provides the person with a behavioral inhibition system (BIS), which is similar to
neuroticism (Carver & White, 1994).
To get an idea for these two neurologically based dimensions of personality, consider your own
reactions to these questionnaire items to assess a tendency toward the BAS:
· When I get something I want, I feel excited and energized.
· When good things happen to me, it affects me strongly.
· I crave excitement and new sensations.
Similarly, consider your own reactions to these questionnaire items to assess a tendency toward
the BIS:
· If I think of something unpleasant is going to happen I usually get pretty “worked up.”
· Criticism or scolding hurts me quite a bit.
· I feel worried when I think I have done poorly at something.
The correlation between people’s scores on the BAS and BIS questionnaires and their prefrontal
lobe asymmetry (as measured by an EEG) is important because the extent of people’s asymmetry

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Individual Brain Structures Involved in Motivation and Emotion 63

corresponds to their typical emotionality (BAS versus BIS; Sutton & Davidson, 1997). People who
score high on BAS items show greater left-side asymmetry, and because of this they show a greater
personality or trait-like sensitivity to reward, eagerness, positive emotion, approach motivation, and
approach-oriented behaviors. People who score high on BIS items show greater right-side asym-
metry, and because of this they show tend to be overly sensitive to punishment, anxiety, negative
emotion, avoidance motivation, and avoidance-oriented behaviors.

Orbitofrontal Cortex
The orbitofrontal cortex lies anatomically beneath the prefrontal cortex, just above the eyes. Anatom-
ically, it is the floor of the prefrontal cortex. It is the cortical brain structure that stores and processes
reward-related information about environmental objects that helps people formulate their preferences
and make their choices between options, such as which product to buy or whether to drink orange
juice or water (Dickinson & Balleine, 2002; O’Doherty, 2004). The orbitofrontal cortex has a direct
connection with the subcortical brain’s reward center (the orbitofrontal-striatal circuit) that allows
it to receive reward-related information from the striatum and, once received, people can remember
the reward value associated with the objects, events, and options they encounter and reencounter.
As we make our way through the day and compare the incentive (or reward) value of the possible
objects and events that might guide our behavior, some objects and some events attract our attention
and serve as attractive incentives to our actions. In a demonstration of the orbitofrontal cortex’s role in
valuing objects, researchers monitored participants’ brains while they looked at a menu and selected
their order. The orbitofrontal cortex is active when people considered their options, remembered what
on the menu was good and what was not, and made their selection among the different environmental
objects (menu items) to pursue (Arana et al., 2003).
The orbitofrontal cortex also inhibits inappropriate actions. It is central to the ability to delay
k gratification, which is essentially quieting the urge for an immediate reward in favor of a more k
advantageous or larger delayed reward. This is a very important capacity, because most long-term
plans involve the ability to put aside those things that are immediately attractive (listen to music, get
something to eat, turn on the TV) in favor of those things that are part of a longer-term strategy to
accomplish goals and complete projects. That is, basic motivations and emotions (e.g., urges, drives,
desires) arise from the subcortical brain and are typically automatic and unconscious—you smell
fresh coffee brewing and you want it. But the orbitofrontal cortex has dense neural connections into
the subcortical brain that allow it to exert self-control (or willpower) over these urges and impulses
for immediate action. It is important to note that this orbitofrontal–subcortical brain communication
system is reciprocal (two-way or bidirectional), because sometimes urges and emotions need to take
precedent over conscious planning, as we do sometimes need to listen to our fears, sense of dis-
gust, and seize on an unexpected opportunity, although we more often need to quiet those urges and
desires to focus our attention and behavior away from such distractions and toward our long-term
goal pursuits.

Ventromedial Prefrontal Cortex

The ventromedial prefrontal cortex groups together a set of interconnected cortical brain areas that
integrate affective-based information from sensory and social cues (Roy, Shohamy, & Wager, 2012).
It represents the affective qualities (or value) of basic sensory rewards, such as tastes, and it is
constantly updating affective representations of internal bodily states. If a stimulus is inherently
appealing (has value), such as a smiling face or a candy bar when one is hungry, extent of ventro-
medial prefrontal cortex activity correlates rather well with how attractive, valuable, or tempting we
find that object to be (Grabenhorst & Rolls, 2011).
In clinical cases in which the person’s ventromedial prefrontal cortex has been damaged (e.g.,
from a stroke), these individuals show emotional impairments and destructive social judgments

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(Damasio, 1994, 1996). This is because the ventromedial prefrontal cortex works for cognitive
valuing and revaluing of emotional inputs that lead to effective decision making (Davidson & Irwin,
1999; Ochsner & Gross, 2005).
The neural connections between the ventromedial prefrontal cortex and subcortical brain are
both dense and bidirectional, just as was the case with the orbitofrontal cortex. This two-way com-
munication allows conscious thought to modulate and control emotion, but it also allows emotion
to inform beliefs, judgments, and decisions. Hence, via its input from the emotion-laden subcortical
brain, the ventromedial prefrontal cortex integrates emotional information with cognitive and social
judgments, including what the person believes to be true (social judgment) and whether the person
believes something is right or wrong (moral judgment) (Cunningham & Zelazo, 2007). The ven-
tromedial prefrontal cortex also receives input from the insula, which can add emotional disgust to
intuitively sway what is believed toward what is not believed. For instance, it is hard to believe that
a food is edible or that a person is trustworthy if either creates a gut feeling of disgust within you.

Dorsolateral Prefrontal Cortex

While the ventromedial prefrontal cortex evaluates the emotional value of basic sensory (unlearned
or natural) rewards, the dorsolateral prefrontal cortex evaluates the learned emotional value of
environmental events and possible courses of action. If someone were to ask you to take a sip of
Coca-Cola and evaluate it emotionally in terms of likes and preferences, you would taste the sensory
properties of the drink in the ventromedial prefrontal cortex, but you would also “taste” and evaluate
the Coca-Cola brand itself in the dorsolateral prefrontal cortex. The idea is that we have a great deal
of learned emotional value and meaning for the objects and events around us, and these emotional
memories are largely stored in the dorsolateral prefrontal cortex. When it comes time to make a
k decision (e.g., What should I buy at the store?), we access this stored environmental information k
from the dorsolateral prefrontal cortex to help us make an emotionally informed decision.
Optimal decision making requires self-control and the dorsolateral prefrontal cortex. The dorso-
lateral prefrontal cortex is involved in the effort to resist temptation during the pursuit of long-term
goals (especially the right dorsolateral prefrontal cortex; Knock & Erst, 2007). Sometimes, we find
ourselves offered an indulgent temptation or a risky opportunity, and neural activity in the dorso-
lateral prefrontal cortex contributes important inhibitory forces during decision making. Basically,
the opposite of urge-based (subcortical) risk-taking is (cortical) self-control. Dorsolateral prefrontal
cortex activations occur (to signal that self-control is occurring) when a person pursues a long-term
reward in favor of a short-term ventral striatum–based reward (McClure, Laibson, Lowenstein, &
Cohen, 2004).
Consider the following experiment that involved people on a diet (Hare, Camerer, & Rangel,
2009). Dieters were shown photographs of various foods, such as an apple and a candy bar. They
were asked to rate how healthy and how tasty each food was for them. When participants rated how
tasty the food was, ventromedial prefrontal cortex activity predicted the “how tasty?” rating. When
participants rated how healthy the food was, dorsolateral prefrontal cortex activity predicted the
“how healthy?” rating. When participants made an actual food choice, the successful dieters showed
strong dorsolateral prefrontal cortex activity, which meant that they exercised self-control and con-
sidered both the food’s taste and its health properties. The unsuccessful dieters did not show this
same dorsolateral prefrontal cortex activity, which shows that they did not exercise self-control and
considered only the food’s rewarding taste. This study showed that strong dorsolateral prefrontal cor-
tex activity dampened down the extent of activity (extent of temptation) participants experienced in
the ventromedial prefrontal cortex (Hare et al., 2009). This is a clear example of what was referred to
earlier in the chapter as bilateral communication (i.e., the dorsolateral prefrontal cortex calms down
ventromedial prefrontal cortex stimulation). Hence, the dorsolateral prefrontal cortex is responsible
for the deployment of self-control.

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Hormones 65

Activity in the dorsolateral prefrontal cortex is also important to keep us from acting selfishly,
because it inhibits our urge for self-interest and therefore contributes positively to harmonious social
interactions (Knock & Erst, 2007). Because of right dorsolateral prefrontal cortex activity, we can
constrain our pursuit of self-interest and make decisions based on our emotional value for social
concerns, such as fairness, equality, and equity (as in the chapter’s opening vignette). Hence, part of
our socioemotional competence lies in right dorsolateral prefrontal cortex activations.

Anterior Cingulate Cortex

There is a lot of conflict in the brain. People wonder “Should I choose this, or should I choose
that? Should I take the immediate reward, or keep working for a later but larger reward? Should I
approach, or should I avoid?” It therefore makes sense that part of the cortical brain exists to make
these judgment calls. That part of the brain is the anterior cingulate cortex, a mid-frontal cortical
brain area that lies above or north of the subcortical brain. It is involved in prioritizing attention,
monitoring conflict, making choices, making decisions, predicting the consequences of actions, and
alerting other brain areas to the need for increased cognitive control to resolve conflict.
In its role as the brain’s information-processing conflict detector, the anterior cingulate cortex
signals and recruits other cortical brain areas to help it to resolve conflict and to exert greater cogni-
tive control. It uses information gained from these other cortical brain structures to select appropriate
action, although it also receives information from the subcortical brain to guide decision making and
action selection (Bush et al., 2002; Matsumoto et al., 2003). The anterior cingulate cortex also evalu-
ates the extent of mental effort required on a task, because cognitive conflict is a good indicator of task
difficulty (Walton, Bannerman, Alterescu, & Rushworth, 2003). Overall, the anterior cingulate cortex
provides “top-down” executive or cognitive control over decision making, goal pursuit, and action.
The core function of the anterior cingulate cortex is to monitor conflicts in information pro-
k cessing and, when conflicts arise, to trigger an increased allocation of cognitive resources, such as k
attention and decision making, to resolve those conflicts in ways that are favorable to one’s goals
(Botvinick, Cohen, & Carter, 2004; Botvinick, Braver, Barch, Carter, & Cohen, 2001; Van Veen,
Cohen, Botvinick, Stenger, & Carter, 2001). For instance, it is in the anterior (and posterior) cin-
gulate cortex where people consciously and deliberatively undertake a cost–benefit analysis as to
whether a goal or a possible course of action has enough reward value associated with it to warrant
an investment of effort (Hayden et al., 2008).

HORMONES
The nervous system uses neurotransmitters (e.g., dopamine) for communication among brain
structures. Alternatively, the endocrine system uses hormones flowing through the bloodstream
to communicate among bodily organs, such as the heart, kidney, and pancreas. While many
hormones are important to motivation and emotion, we highlight these three: cortisol, oxytocin, and
testosterone. Other hormones are integral to basic biological motivations such as hunger and thirst,
and these hormones will be featured in Chapter 4.

Cortisol
Cortisol is the so-called stress hormone. Cortisol is a hormonal product of the reactivity of
the hypothalamic–pituitary–adrenocortical system (Dickerson & Kemeny, 2004; Stansbury &
Gunnar, 1994; Susman, 2006). In response to stressful events, subcortical brain structures (e.g., the
amygdala) stimulate the hypothalamus to stimulate the pituitary gland, which leads the adrenal
gland to increase cortisol production and release it into the bloodstream. Cortisol activation
occurs during social-evaluative threats (e.g., public speaking), relationship conflict (Powers,
Pietromonaco, Gunlicks, & Sayer, 2006), and being interpersonally controlled (Reeve & Tseng,
2011) or devalued/rejected (Stroud, Salovey, & Epel, 2002). Cortisol deactivation (destress) occurs

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66 Chapter 3 The Motivated and Emotional Brain

during social support (Kirschbaum, Klauer, Filipp, & Hellhammer, 1995; Reeve & Tseng, 2011;
Taylor et al., 2010).
Generally speaking, cortisol reactivity serves a short-term adaptive function, as it mobilizes
attention and energy in response to a social evaluative threat (Dickerson & Kemeny, 2004). Longer-
term, however, chronic cortical reactivity (repeated hypothalamic–pituitary–adrenocortical system
activation, as from long-term exposure to a conflictual relationship) takes a cumulative toll on the
body, a phenomenon termed “allostatic load” (McEwen, 1998). Cortisol-induced allostatic load puts
the individual at risk of negative biological outcomes such as diabetes and hypertension, but it has
further been linked to maladaptive cognitive outcomes, such as poor memory, impaired problem
solving, and poor intellectual functioning (Brown & Suppes, 1998; Kirschbaum et al., 1996).

Oxytocin
Oxytocin is known as the bonding hormone. Oxytocin release supports the “tend and befriend stress
response” that helps explain why people seek counsel and confide in friends during the stress-
ful events in their lives. Oxytocin increases the salience or attention-getting qualities of social-
interpersonal cues, such as emotion recognition and empathy (Bartz, Zaki, Bolger, & Ochsner, 2011),
and it is associated with social engagement and the seeking of sociability that can calm and suppress
arousal, stress, and depression (Heinrichs, Bumgartner, Kirschbaum, & Ehlert, 2003).
Oxytocin also raises levels of trust in others (Campbell, 2010; Kosfeld et al., 2005). Oxytocin
supports the formation and maintenance of attachment bonds among people (Lim & Young, 2006),
and it boosts social behaviors such as trusting others, being generous, sharing resources, and being
cooperative (Kosfeld et al., 2005; Zak et al., 2007). The tend-and-befriend coping response can
be highly effective during times of stress (compared to the well-known “fight-or-flight” coping
response) as people seek the counsel, support, and nurturance of others. It is the hormone of
k oxytocin that underlies this tend-and-befriend coping response during times of stress, and this k
hormone also supports attachments and bonds with those who are befriended (Feldman, Weller,
Zagoory-Sharon, & Levine, 2007; Pedersen, 2006).
One interesting feature of oxytocin is that it can be delivered to a person via the squirt of a nasal
spray. In experiments in which people in the experimental condition received an intranasal oxytocin
administration while people in the control condition receive only a placebo, those in the experimen-
tal group showed significant increases in postspray trust, cooperation, eye contact, empathy, and
social bonding (Declerck, Boone, & Kiyonari, 2010; Guastella, Mitchell, & Dadds, 2008; Kosfeld,
Heinrichs, Zak, Fischbacher, & Fehr, 2005; Van IJzendoorn & Bakermans-Kranenburg, 2012).

Testosterone
The steroid hormone of testosterone is associated with high competition, status-seeking, and sexual
motivation (Bancroft, 2002). High testosterone encourages competition. For instance, high testos-
terone levels help Wall Street stockbrokers make more money (compete better) during the day’s
trading. More specifically, high testosterone is associated with status-seeking behavior (winning a
competition), and it is most strongly associated with status-seeking behaviors after social status has
been questioned or threatened (after losing a battle or competition for social dominance) (Josephs,
Newman, Brown, & Beer, 2003; Josephs, Sellers, Newman, & Mehta, 2006). Testosterone also
underlies the mating effort—the investment of time and energy into same-sex competition and
mate-seeking behavior (Ellison, 2001). Unmarried men, for instance, have higher testosterone levels
than do married men (Gray et al., 2004); men who are not in a committed relationship have higher
testosterone levels than do men who are in a committed relationship (Burnham et al., 2003); and
men who are not fathers have higher testosterone than do men who are fathers (Gray et al., 2002).
High levels of testosterone are also associated with having affairs, while low levels are associated
with better parenting (e.g., higher nurturance).

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Summary 67

SUMMARY
When thinking about the brain, most people focus on its cognitive and intellectual functions, includ-
ing thinking, learning, and problem-solving. But it is more—it is also a motivated and emotional
brain. It generates wants, appetites, urges, needs, reward, cravings, desires, pleasure, feelings, mood,
fear, anxiety, anger, and the full range of the emotions. The neuroscience of motivation and emo-
tion focus on understanding how environmental objects and day-to-day event activate specific brain
structures and how these brain structures, in turn, are associated with the motivational and emotional
states that energize, direct, and sustain behavior.
At a general level, the motivated and emotional brain consists of an outer cortical brain and an
inner subcortical brain. The subcortical brain is associated with basic urges and impulses and with
emotion-rich motivations such as hunger, thirst, anger, fear, anxiety, pleasure, desire, reward, and
wanting. These urges and emotions are largely unconscious, automatic, and impulsive. The corti-
cal brain is associated with cognitively rich motivations such as goals, plans, strategies, values, and
beliefs about the self. The mental events are largely conscious, deliberate, and revolve around cogni-
tive or executive control. Both the cortical and subcortical brain features many individual structures,
but it is important to note that these individual structures are linked together by a network of neural
superhighways to communicate reciprocally with each other. The overall picture of brain function
is therefore not one in which individual brain structures are associated with individual functions
(e.g., the amygdala does this, the nucleus accumbens does that) but, rather, one of interconnectivity
and the brain working in a highly integrated way.
Nine subcortical brain structures are closely involved in motivation and emotional states.
The reticular formation regulates arousal, alertness, and the neural process of awakening the brain’s
motivational and emotional concerns. The amygdala detects, learns about, and responds to the
stimulus properties of environmental objects, including both threat-eliciting and reward-eliciting
k associations. The basal ganglia (caudate nucleus, putamen, substantial nigra, and globus pallidus) k
contribute to the motivational invigoration and inhibition of movement and action. The ventral
striatum (nucleus accumbens) and ventral tegmental area constitute the brain’s reward center.
The ventral tegmental area manufactures and releases dopamine that is received by the nucleus
accumbens to produce pleasure and liking. The hypothalamus is responsive to natural rewards in the
regulation of eating, drinking, and mating, and it also regulates both the endocrine and autonomic
nervous systems.
Six cortical brain structures are closely involved in motivation and emotional states. The
insula monitors bodily states to produce both positive and negative gut-felt feelings, and it also
processes feelings associated with risk, uncertainty, intrinsic motivation, empathy, and personal
agency. The prefrontal cortex is involved in making plans, setting goals, formulating intentions.
Right hemispheric activity is associated with negative affect and “no-go” avoidance motivation,
while left hemispheric activity is associated with positive affect and “go” approach motivation. The
orbitofrontal cortex stores and processes reward-related values of environmental objects and events
to formulate preferences and make choices between options. The ventromedial prefrontal cortex
evaluates the unlearned emotional value of basic sensory rewards and internal bodily states and is
responsible for emotional control. The dorsolateral prefrontal cortex evaluates the learned emotional
value of environmental events and possible courses of action, and it is responsible for control
over urges and risks during the pursuit of long-term goals. The anterior cingulate cortex monitors
motivational conflicts. It resolves those conflicts by recruiting other cortical brain structures to exert
cognitive control over basic urges and emotions.
While the nervous system relies on neurotransmitters to communicate among brain structures,
the endocrine system relies on hormones flowing through the bloodstream to communicate between
bodily organs, such as the heart. While many hormones are important for motivation and emotion,
cortisol, oxytocin, and testosterone are particularly important. Cortisol produces an energized

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68 Chapter 3 The Motivated and Emotional Brain

stress response when the person is exposed to a social-evaluative threat, such as public speaking or
relationship conflict. Oxytocin produces an affiliation-based tend-and-befriend stress response when
people seek counsel and confide in friends during the stressful events in their lives. Testosterone
produces competitive status-seeking behaviors, such as when a man’s social status has been
questioned or threatened.

READINGS FOR FURTHER STUDY

Subcortical Brain
Berridge, K. C. (2004). Motivational concepts in behavioral neuroscience. Physiology and Behavior, 81, 179–209.
Kuhnen, C. M., & Knutson, B. (2005). The neural basis of financial risk taking. Neuron, 47, 763–770.
McClure, S. M., Laibson, D. I., Loewenstein, G., & Cohen, J. D. (2004). Separate neural system value immediate and
delayed monetary rewards. Science, 506, 503–507.
Pessiglione, M., Schmidt, L., Draganski, B., Kalisch, R., Lau, H., Dolan, R. J., & Frith, C. D. (2007). How the brain
translates money into force: A neuroimaging study of subliminal motivation. Science, 316, 904–906.

Cortical Brain
Becharea, A., Damasio, H., Tranel, D., & Damasio, A. R. (1996). Deciding advantageously before knowing the advanta-
geous strategy. Science, 275, 1293–1295.
Craig, A. D. (2009). How do you feel—now? The anterior insula and human awareness. Nature Review: Neuroscience, 10,
59–70.
Sutton, S. K., & Davidson, R. J. (1997). Prefrontal brain asymmetry: A biological substrate of the behavioral approach and
inhibition systems. Psychological Science, 8, 204–210.
Hare, T. A., Camerer, C. F., & Rangel, A. (2009). Self-control in decision-making involves modulation of the vmPFC
valuation system. Science, 324, 646–648.

Hormones
Dickerson, S. S., & Kemeny, M. E. (2004). Acute stressors and cortisol responses: A theoretical integration and synthesis
k k
of laboratory research. Psychological Bulletin, 130, 355–391.
Heinrichs, M., Baumgartner, T., Kirschbaum, C., & Ehlert, U. (2003). Social support and oxytocin interact to suppress
cortisol and subjective responses to psychosocial stress. Biological Psychiatry, 54, 1389–1398.