Psychologia, 2011, 54, 222–233

BRAIN MECHANISMS OF HAPPINESS

Shintaro FUNAHASHI

Kokoro Research Center, Kyoto University, Japan.

Happiness is a kind of subjective feeling of positive emotions. Since the

psychological process of reward is closely linked to pleasure, studies to elucidate the
brain mechanisms of happiness have focused on the neural circuitry that processes
reward information. Brain imaging studies have revealed that the orbitofrontal
cortex, cingulate cortex, medial prefrontal cortex, insula, nucleus accumbens,
ventral pallidum, substantia nigra, and ventral tegmental area are the key areas
related to happiness. When we consider the brain mechanisms of happiness, we
need to distinguish between the neural mechanisms for liking and wanting, and
between those that encode and cause pleasant feelings. Among these brain areas,
subcortical structures participate in “wanting”, while cortical areas participate in
“liking”. Orbitofrontal activity has been shown to encode the subjective experience
of pleasure. However, it is not clear whether orbitofrontal activity actually causes
pleasant feelings. Since the neuroscience of happiness is still in the primitive stage,
further experiments are needed to elucidate the brain mechanisms of happiness.

Key words: reward, dopamine, orbitofrontal cortex, ventral striatum, liking,

wanting

Happiness is a kind of subjective feeling of positive emotions. Happiness is closely

linked to pleasure and pleasant feelings. The word “happiness” is explained in the Oxford
English Dictionary as (1) good fortune or luck in life or in a particular affair (success,
prosperity), (2) the state of pleasurable content of mind, which results from success or the
attainment of what is considered good, and (3) successful or felicitous aptitude, fitness,
suitability, or appropriateness. In the Random House Dictionary for the English
Language, the word “happiness” is defined as “results from the possession or attainment
of what one considers good,” and synonyms include pleasure, joy, exhilaration, bliss,
contentedness, delight, enjoyment, and satisfaction. Thus, we can experience happy
feelings at different times and in a variety of places, situations, and contexts.
Since the psychological process of reward is closely linked to pleasure or liking,
psychological and neuroscience studies have focused on the neural circuitry that processes
reward information to understand how pleasure or happy feelings are generated in the
brain and which brain structures participate in generating pleasure and happiness.
Although the neuroscience of happiness is still in the primitive stage compared with the
neuroscience of perception and motor control, key brain structures or hotspots for pleasure
have been clarified using functional brain imaging techniques (e.g., positron emission
tomography (PET) and functional magnetic resonance imaging (fMRI)). In this article, I
Correspondence concerning this article should be addressed to Shintaro Funahashi, Kokoro Research
Center, Kyoto University, 46 Yoshida-Shimoadachi, Sakyo-ku, Kyoto 606-8501 Japan (e-mail:
funahashi.shintaro.2z@kyoto-u.ac.jp).

222
 BRAIN MECHANISMS OF HAPPINESS 223

will first consider brain structures that presumably participate in generating feelings of
pleasure. Next, I will discuss several important brain areas, including the basal ganglia,
the orbitofrontal cortex, the anterior cingulate cortex, and the insula, to consider the neural
mechanisms of happiness.

NEUROANATOMY OF HAPPINESS

Recent advances in affective neuroscience research have revealed brain networks

related to the reward system and neurotransmitters related to pleasure and positive
emotions. The brain areas that are activated coincident with pleasure and positive
emotions are widespread. These areas include the orbitofrontal cortex, the cingulate
cortex, the medial prefrontal cortex, the insula, the nucleus accumbens, the ventral
pallidum, the substantia nigra, and the ventral tegmental area. These brain areas are often
called “hedonic hotspots” (Kringelbach & Berridge, 2009). The pleasure-generating
ability of these brain areas has been demonstrated by the electrical stimulation of these
structures or the direct application of neurochemicals, such as amphetamine, to these
structures. These procedures usually produce a doubling or more of the number of
responses than are normally elicited by a natural reward such as sucrose. Therefore, these
observations suggest that either the electrical or chemical stimulation of these brain
structures can generate pleasure or positive emotions.
A reward activates the reward system or hedonic hotspots and generates feelings of
pleasure or positive emotions (e.g., liking). At the same time, a reward also activates
motivational systems and produces incentive behavior (e.g., wanting). For example, when
rats accidentally press a key and receive electrical stimulation through an electrode
implanted in the hypothalamus, they try to press the key as many times as they can to get
more electrical stimulations, which is the famous “brain stimulation reward (BSR).” In
this situation, the electrical stimulation in the hypothalamus acts as a positive
reinforcement. The rats seem to really want to receive the stimulation. However, the
mechanism related to “wanting” is not always linked to that for “liking.” Thus, while it is
clear that rats apparently want to receive electrical stimulations by pressing the key, it is
not clear whether they are really gaining pleasurable feelings or positive emotions by
electrical stimulation at the hypothalamus. The discrepancy between “wanting” and
“liking” can be seen in addictive behavior. While addicts compulsively want to take
drugs, they do not always want to do so at a conscious level. They sometime have
negative emotions and do not like them.
The electrical stimulation of subcortical structures, such as the nucleus accumbens,
the lateral hypothalamus, and the ventral tegmental area, has been known to produce
strong incentive behaviors in animals. Therefore, subcortical structures are considered to
participate in the mechanisms of “wanting.” In addition, since these structures belong to
the mesolimbic dopamine system, this system is considered to play an important role in
the mechanism of “wanting”.
On the other hand, cortical hedonic hotspots including the orbitofrontal cortex, the
224 FUNAHASHI

insula, and the medial prefrontal cortex are considered to participate in liking and
pleasure. These brain areas have anatomical connections to subcortical hedonic hotspots.
In addition, neuroimaging studies have shown that these brain areas encode hedonic
evaluations including the anticipation, experience, and memory of pleasant stimuli.
Kringelbach & Berridge (2009) pointed out the importance of the distinction between
brain activities that “encoded” pleasure and that “caused” pleasure. According to their
definition, neural “coding” can be inferred by measuring brain activity correlated to a
pleasant stimulus, whereas causation is inferred on the basis of a change in pleasure as a
consequence of a brain manipulation, such as a lesion or stimulation. The distinction
between “encoding” and “causing” pleasure is important when we consider how a
particular cortical area participates in neural mechanisms of happiness.

PLEASURE CENTER IN THE BRAIN

If a pleasure center exists somewhere in the brain, and if we wish to understand the
nature of this pleasure center, we need to consider the distinction between liking and
wanting and the distinction between brain activity that encodes pleasure and that which
causes pleasure. However, since encoding and causing are often associated with the same
treatment, it is not always easy to make such a distinction. In this section, we will look at
some animal experiments that were performed to identify hedonic hotspots in the brain.

Brain Stimulation Reward (BSR)

In the early 1950s, Olds and Milner discovered that a rat continued to perform key-
pressing responses in an operant box when electrical stimulation was applied through an
electrode implanted within the lateral hypothalamus when the rat pressed a key (Olds &
Milner, 1954; Olds, 1958). Subsequent studies showed that electrical stimulation in the
lateral hypothalamus as well as regions of the medial forebrain bundle produced a very
powerful reward effect (Crow, 1972). Rats continuously pressed the key several hundred
times per hour under electrical stimulation of the lateral hypothalamus or regions of the
medial forebrain bundle. When a rat had unlimited access to a key to receive electrical
stimulation at the lateral hypothalamus, amazingly, the rat continued to press the key for
several days, without rest and without taking food or water (Olds, 1958). Thus, electrical
stimulation at the lateral hypothalamus produces a very powerful reward effect and can be
used as very powerful positive reinforcement. Therefore, this stimulation has been called
the brain stimulation reward (BSR) (Olds, 1962, 1976).
To prove that the BSR observed in rat experiments is linked to a positive emotion or
pleasure, the effect of the BSR was compared with the effect of a painful electric foot-
shock (Olds, 1962). As a result, rats accepted strong electric foot-shocks to get the BSR,
even though the electric foot-shock was stronger than that which the rats would not accept
to get food or water. Since an electric foot-shock is a painful stimulus for the rat, it should
generate strong negative emotions (e.g., fear). However, the observation that rats chose to
receive the BSR despite painful foot-shocks reveals that the BSR can produce a stronger
 BRAIN MECHANISMS OF HAPPINESS 225

positive emotion or stronger pleasant feeling in rats to overcome an unpleasant negative

emotion. Based on these observations, the lateral hypothalamus has been referred to as a
“pleasure center” in the brain.
While Olds found that the lateral hypothalamus was a pleasure center in the brain,
subsequent studies showed that several other brain areas also exhibit a BSR. The brain
structures along the medial forebrain bundle are the major regions that are related to the
BSR (Crow, 1972). These include the lateral hypothalamus, the ventral tegmental area,
the substantia nigra, and the septum. The BSR effect is also produced by stimulation of
the cerebral cortex, including the medial and orbital prefrontal cortex, and the cingulate
cortex (Rolls, 1974). Limbic areas including the hippocampus and the amygdala also
produce a BSR (Rolls, 1974).
Although the BSR effect was initially observed in rats and has been extensively
studied using rats, electrical stimulation of these brain structures has also been shown to
produce a BSR effect in monkeys (Rolls et al., 1980) and humans (Delgado, 1976). Heath
(1972) set a stimulation electrode within the septal area in the human brain and found that
human patients reported a pleasant feeling when they were stimulated through the
implanted electrode and that, when given access to a button, they repeatedly pressed the
button to stimulate their own brain. Thus, these results indicate that our brain includes
structures that produce pleasant feelings under electrical stimulation. Hence, by
stimulating these structures, we can elicit pleasant and happy feelings. Therefore, the
brain structures that produce a BSR effect should be hedonic hot spots.

Dopamine and Pleasant Feelings

Behavioral studies have established that brain structures along the medial forebrain
bundle produce the BSR effect. Next, we wish to consider which neurotransmitters or
neurochemicals play a role in this BSR effect. Pharmacological studies have revealed that
dopamine is a neurotransmitter that contributes to the BSR effect (Phillips et al., 1989;
Wise & Rompre, 1989; Garis et al., 1999; Spanagel & Weiss, 1999). It has been shown
that the administration of dopamine agonists or antagonists modulated the BSR effect.
For example, the microinjection of a dopamine antagonist (e.g., amphetamine) into the
lateral hypothalamus attenuated the BSR effect in rats (Esposito et al., 1980). Careful
mapping studies showed that the brain structures associated with the BSR effect
correspond to the structures where dopamine neurons exist or dopamine fibers are present
or terminate (Crow, 1976; Wise, 1981; Prado-Alcala & Wise, 1984; Prado-Alcala et al.,
1984). Especially, it has been shown that the medial forebrain bundle includes massive
dopamine fibers. Both the meso-cortical dopamine fibers and the nigrostriatal dopamine
fibers pass through the lateral hypothalamus, which is a part of the medial forebrain
bundle and the most effective brain area of the BSR. Therefore, electrical stimulation
through electrodes implanted in the lateral hypothalamus or the medial forebrain bundle
could directly activate dopamine fibers passing these structures (German & Bowden,
1974; Wise & Rompre, 1989). Thus, activation of the dopamine system could produce the
BSR effect, and hence generate pleasant and happy feeling in animals and humans.
226 FUNAHASHI

Basal Ganglia
In the basal ganglia, the nucleus accumbens and the ventral pallidum have been
considered to participate in the mechanisms of liking that cause pleasure responses. These
brain areas possibly enhance liking responses to sensory stimuli that produce pleasant
feelings such as a sweet taste or opioid stimulus. Berridge (1996) stated in his review that
a reward contains distinguishable functional components, i.e., liking (pleasure/
palatability) and wanting (appetite/incentive motivation), and that liking is mediate by
neurotransmitter systems such as opioid systems and anatomical structures such as the
ventral pallidum, while the mediation of wanting involves mesocortical dopamine systems
and a division of the nucleus accumbens. For example, microinjection of a µ-opioid
agonist in either the nucleus accumbens or the ventral pallidum of rats enhanced positive
orofacial reactions elicited by the hedonic effect of a sucrose solution (e.g., tongue
protrusions), which are homologous to affective facial expressions of liking in primates
and human infants (Smith & Berridge, 2005). Opioid stimulation in the nucleus
accumbens and the ventral pallidum also increased motivational wanting behavior for
food (e.g., increased eating behavior), but did not increase liking responses to taste stimuli
(Smith & Berridge, 2005). Thus, opioid stimulation in the nucleus accumbens or the
ventral pallidum enhances hedonic liking responses to a sweet taste and appetitive
wanting responses to a food reward. Subsequently, Smith & Berridge (2007) showed that
the nucleus accumbens and the ventral pallidum were both needed to enhance liking
responses and cooperate as a single hedonic circuitry. However, the nucleus accumbens
dominated wanting responses (e.g., eating and food intake), independent of activation of
the ventral pallidum.
The evidence that the ventral pallidum mediates liking responses is supported by
observations that liking responses are abolished in rats with lesions in the ventral
pallidum. Cromwell & Berridge (1993) made excitotoxic lesions in the bilateral ventral
pallidum and examined the effect of these lesions on affective reactions to the application
of sucrose solutions into the mouth. They found the enhancement of aversive or disliking
responses (gapes, chin-rubbing, head-shaking, and forelimb flails) to sucrose application.
Miller et al. (2006) reported a human patient (Mr. A) who had a selective bilateral lesion
of the globus pallidus. Mr. A, a 34-year-old man, was a heavy drug user. He was brought
to an emergency room after consuming cocaine, Ecstasy (MDMA), oxycodone, and
methadone at a party. After this overdose, he became depressed and complained of a
depressed mood, anhedonia, low energy, difficulties concentrating and remembering,
feelings of hopelessness and guilt, poor self-esteem, and social isolation. He reported that
he no longer experienced pleasure from drinking alcohol. An MRI scan of his brain
revealed a selective bilateral lesion of the globus pallidus.
Thus, the ventral pallidum and the nucleus accumbens seem to participate in liking
mechanisms that cause pleasure responses. Therefore, these brain structures can be
considered subcortical pleasure centers or hot spots and are important when considering
the neurobiology of happiness. Experimental results suggest that the ventral pallidum
participates in enhancing liking responses, while the nucleus accumbens participates in
enhancing wanting responses. However, further studies are needed before we can make
 BRAIN MECHANISMS OF HAPPINESS 227

firm conclusions regarding how these structures contribute to the generation of happy
feelings.

CORTICAL HOT SPOTS FOR PLEASURE

Orbitofrontal Cortex
The orbitofrontal cortex contains the taste area (Ongur & Price, 2000). This taste
area has been shown to represent the reward value of taste, in that neurons responded to
food only when the animal was hungry (Rolls, 2000). The orbitofrontal cortex also
contains the olfactory area (Ongur & Price, 2000). This olfactory area represents the
reward value of odor, and olfactory neurons in this area responded to the smell of a food
only when the animal was hungry (Critchley & Rolls, 1996). With regard to orbitofrontal
neurons, it has been shown that neuronal responses to a particular food decrease when the
animal is fed this food to satiety. This phenomenon is known as “sensory-specific satiety.”
Thus, single-neuron studies have indicated that orbitofrontal neurons represent the reward
value of food and change their activities depending on the relative reward value.
However, these studies have provided no evidence indicating that brain activity in the
orbitofrontal cortex is correlated with a subjective report of pleasure or pleasant feelings.
Sensory-specific satiety is useful for studying affective representations in the brain, since
it provides means of altering the affective value of a stimulus without modifying its
physical nature. Therefore, Kringelbach et al. (2003) used the phenomenon of sensory-
specific satiety and examined brain activity using fMRI. In this experiment, the human
subjects were given either chocolate milk, tomato juice, or a tasteless control solution and
asked to rate both pleasantness and intensity. They found a significant correlation
between the activation of the orbitofrontal cortex and the decrease in subjective
pleasantness when a particular liquid was taken to satiety. In addition, the decrease in
orbitofrontal activation was specific to the particular liquid taken by the subject, which
constitutes evidence for a neural correlate of sensory-specific satiety in humans. Thus,
these studies indicate that the orbitofrontal cortex represents not only the reward value of
food but also the subjective pleasantness when the food is consumed.
Several recent studies have shown that the encoding of pleasure reaches the
orbitofrontal cortex. Neuroimaging studies have revealed correlations between brain
activation and subjective pleasantness in human subjects. Blood & Zatorre (2001)
examined the correlation between the magnitude of changes in cerebral blood flow and the
magnitude of subjective pleasant emotional responses to subject-selected music. They
also confirmed subjective reports of the subjects’ emotions by measuring changes in heart
rate, electromyogram, and respiration. Correlated cerebral blood flow changes were
observed in the orbitofrontal cortex, in addition to the ventral striatum, the amygdala, and
the ventromedial prefrontal cortex. Vollm et al. (2004) used highly addictive drugs,
amphetamines, to drug naïve human subjects and examined correlations between brain
activation measured by fMRI and subjective ratings of “mind-racing,” which was defined
as “a state where your thoughts run far too quickly as if they were racing. One thought is
228 FUNAHASHI

replaced by another very quickly and it is difficult to hold on to your thought.” They
observed activation of the medial orbitofrontal cortex under the administration of
amphetamines and a significant correlation between activation of the orbitofrontal cortex
and ratings of the mind-racing experience. In addition, Small et al. (2001) used chocolate
and measured regional cerebral blood flow using PET. After they ate a piece of chocolate,
subjects were asked to rate how pleasant or unpleasant the chocolate was and how much
they did or did not want another piece of chocolate. They found that the caudomedial
orbitofrontal cortex was active when subjects were highly motivated to eat chocolate and
rated the chocolate as very pleasant, while the caudolateral orbitofrontal cortex was active
when subjects ate chocolate despite being satiated. Based on these results, they concluded
that there is a functional segregation of the neural representation of pleasant and
unpleasant emotion within the orbitofrontal cortex.
Interestingly, the magnitude of the activity of the orbitofrontal cortex tracks changes
in the strength of subjective pleasantness. For example, the magnitude of the brain
activity was reduced when the reward value of one food was reduced by eating it to
satiety. However, when another new food was introduced as a reward, the magnitude of
the activity recovered to a high level (e.g., Kringelbach et al., 2003). Thus, the
orbitofrontal cortex is an important candidate for coding of the subjective experience of
pleasure or happiness.
The orbitofrontal cortex is an important brain region for processing pleasantness and
unpleasantness. Kringelbach & Rolls (2004) performed a meta-analysis of a large number
of neuroimaging studies and proposed two distinct trends of neural processing in the
orbitofrontal cortex. One trend is a mediolateral distinction. The medial orbitofrontal
cortex is related to positive emotion. Medial orbitofrontal activity has been shown to
reflect the reward value of many different food or liquid rewards. The medial
orbitofrontal cortex was activated in relation to the pleasantness of stimuli, such as
whether the taste and smell of stimuli are pleasant (Rolls et al., 2003a) and the degree of
this pleasantness (De Araujo et al., 2003). On the other hand, the lateral orbitofrontal
cortex is related to negative emotion. Lateral orbitofrontal activity is known to reflect the
unpleasantness of stimuli, such as monetary loss (O’Doherty et al., 2001) or painful
touching (Rolls et al., 2003b). Since such a negative emotion would be important for
changing ongoing behavior, lateral orbitofrontal activity may not be directly related to the
negative emotion, such as unpleasantness, but may encode a signal to escape from the
stimulus. The second trend is a posterior-anterior distinction. It has been shown that an
abstract-concrete gradient is present along the posterior-anterior axis. More complex or
abstract reinforcers (e.g., the gain or loss of money) are represented more anteriorly in the
orbitofrontal cortex (O’Doherty et al., 2001), while simpler and concrete reinforcers (e.g.,
a good taste or pain) are represented more posteriorly (Kringelbach et al., 2003). Thus,
within the orbitofrontal cortex, the medial-lateral hedonic gradient interacts with the
posterior-anterior abstract-concrete gradient.
Although there is some evidence that orbitofrontal activity encodes the subjective
experience of pleasure or happiness, it is not yet clear whether orbitofrontal activity
actually causes positive emotion or pleasure. However, it has been shown that damage to
 BRAIN MECHANISMS OF HAPPINESS 229

the orbitofrontal cortex impairs pleasure-related decision-making. Baxter et al. (2000)

showed that rhesus monkeys with a surgical disconnection between the orbitofrontal
cortex and the amygdala were unable to adjust their choice behavior during performance
of a visual discrimination task after a change in the outcome (different food pellets).
However, this lesion did not affect their motivation to work for a food reinforcer or food
preferences. The orbitofrontal cortex has been shown to use learned representations of
outcomes to guide behavior. Pickens et al. (2003) first trained rats to associate a light
conditioned stimulus with a food outcome, and then devalued the food by pairing it with
an unpleasant stimulus. They then examined responses to the conditioned stimulus. They
found that the response to the conditioned stimulus was attenuated in rats with
orbitofrontal lesions when the lesion was made after light-food conditioning. Therefore,
they concluded that the orbitofrontal cortex seems to be important for maintaining these
stimulus-outcome representations in memory, updating these representations with new
information, or using these representations for behavior. Thus, the orbitofrontal cortex
seems to participate in pleasure-related decision-making. However, since patients with
damage to the orbitofrontal cortex still exhibit normal pleasure responses (Anderson et al.,
1999; Hornak et al., 2003), some caution is required when we consider whether the
orbitofrontal cortex generates positive emotion or pleasure.

Anterior insula
Almost all recent imaging studies of emotion show activation of the anterior insula
(AI) in human subjects experiencing a variety of emotional feelings, including maternal
and romantic love, anger, fear, sadness, happiness, sexual arousal, disgust, aversion,
unfairness, inequity, indignation, uncertainty, disbelief, trust, empathy, and beauty (Craig,
2009). For example, activation of the left AI was reported when mothers viewed photos
of their own children (Leibenluft et al., 2004). Activation of the left AI was reported
while subjects were either seeing or making a smile (Jabbi et al., 2007). Activation in the
AI was found when subjects listened to happy voices (Johnstone et al., 2006), and has
been associated with hearing pleasant music (Koelsch et al., 2006). Further, selective
activation of the left AI was found when subjects were experiencing joy (Takahashi et al.,
2008). Thus, the AI apparently is activated in association with a variety of subjective
feelings including happiness or pleasant feelings.

Medial Prefrontal Cortex

Recently, the medial prefrontal cortex including the anterior cingulate cortex has
been considered to be an important brain structure for social cognition processes, such as
self-reflection, person perception, and making inferences about other’s thoughts (Amodio
& Frith, 2006). However, the medial frontal cortex is not a uniform area. This area has
been divided into several functionally different sub-regions (Beckmann et al., 2009).
Functional brain imaging studies show that the anterior dorsal cingulate sulcus is active
when the link between an action and reinforcement is critical, especially during error
detection, or when action selection is guided by the expectation of a reward (Rushworth et
al., 2007). This region is also important for detecting when actions are potentially in
230 FUNAHASHI

conflict (Botvinick et al., 2004). In contrast, the ventral cingulate region is active when
subjects experience pain or during autonomic arousal (Vogt, 2005). Importantly, both
ventral and anterior dorsal cingulate regions are active during emotion and social
interaction (Amodio & Frith, 2006). Ochsner et al. (2004) examined neural activity while
subjects viewed pictures depicting a person in a positive, negative, or neutral scene. For
each trial, subjects were asked to judge their own affective response, the affective
response of the person in the picture, or whether the picture depicted an indoor or outdoor
scene. Anterior cingulate activation was observed when subjects judged their own
affective responses. Thus, the anterior cingulate area could participate in the self-
knowledge of the subject’s subjective feelings. Cingulate participation in the self-
knowledge of one’s own subjective feeling could be important for us to experience happy
feelings.

CONCLUSIONS

Happiness is a kind of subjective feeling of positive emotions. Since the

psychological process of reward is closely linked to pleasure, studies to elucidate the
neural mechanisms of happiness have focused on the neural circuitry for processing
reward information. Although the neuroscience of happiness is still in the primitive stage
compared with the neuroscience of perception and motor control, key brain areas related
to happiness have been clarified using functional brain imaging techniques. These brain
areas include the orbitofrontal cortex, the cingulate cortex, the medial prefrontal cortex,
the insula, the nucleus accumbens, the ventral pallidum, the substantia nigra, and the
ventral tegmental area (Table 1).

Table 1. Brain Areas Related to Brain Stimulation Reward, Wanting, and/or Liking.

Brain stimulation reward Wanting Liking

Cerebral cortex
Medial orbitofrontal cortex ○ ○
Cingulate cortex ○ ○
Anterior insula ○ ○
Subcortical structures
Nucleus accumbens ○ ○ ○
Ventral pallidum ○ ○
Substantia nigra ○ ○
Ventral tegmental area ○ ○
Septum ○ ○
Lateral hypothalamus ○ ○
 BRAIN MECHANISMS OF HAPPINESS 231

When we consider neural mechanisms of happiness, we need to distinguish between

neural mechanisms for liking and those for wanting, and between those that encode and
those that cause pleasant feelings. A reward activates hedonic hot spots and generates
pleasant feelings or positive affections (liking). At the same time, the reward also
activates motivational systems and produces incentive behavior (wanting). Electrical
stimulation of the nucleus accumbens, the lateral hypothalamus, and the ventral tegmental
area has been shown to produce strong incentive behaviors in animals. Therefore, these
subcortical structures are considered to participate in “wanting” mechanisms. On the
other hand, cortical areas, including the orbitofrontal cortex, the insula, and the medial
prefrontal cortex, are considered to participate in “liking” mechanisms.
The neural mechanism for “encoding” pleasant feelings is inferred by measuring
brain activity correlated to a pleasant stimulus, whereas the neural mechanism that
“causes” pleasant feelings is inferred on the basis of a change in pleasure as a consequence
of a brain manipulation, such as a lesion or stimulation. Since encoding and causing often
go together for the same treatment, it is not always easy to distinguish between the two.
There is some evidence that orbitofrontal activity encodes the subjective experience of
pleasure or happiness. However, it is not yet clear whether orbitofrontal activity actually
causes positive emotion or pleasure. Further experiments are needed to identify the brain
area that causes the subjective experience of happiness.

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(Manuscript received 1 September, 2011; Revision accepted 20 October, 2011)