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Discov Med. 2010 June ; 9(49): 579–587.

The Functional Neuroanatomy of Pleasure and Happiness

Morten L. Kringelbach, D.Phil. and

Department of Psychiatry, Warneford Hospital, University of Oxford, Oxford, United Kingdom and
Centre for Functionally Integrative Neuroscience (CFIN), University of Aarhus, Aarhus, Denmark
Kent C. Berridge, Ph.D.
Department of Psychology, University of Michigan, Ann Arbor, Michigan 48109, USA

Abstract
Over fifty years ago the discovery that rats would work to electrically stimulate their brains
suggested the intriguing possibility that bliss could be achieved through the use of ‘pleasure
electrodes’ implanted deep within the brain. Subsequent research has failed to bring about this
brave new world of boundless pleasure, but more recent findings have started to throw new light
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on the intriguing links between brain mechanisms of pleasure and happiness. We discuss these
findings of the underlying neural mechanisms and functional neuroanatomy of pleasure in the
brain. In particular we address how they may come to shed light on our understanding of the brain
basis of happiness. Beyond sensory pleasures, we examine how higher pleasures may be related to
the brain’s default networks, especially in orchestrating cognitive aspects of the meaningfulness
important to happiness. We also address how understanding of the hedonic brain might help
alleviate the suffering caused by the lack of pleasure, anhedonia, which is a central feature of
affective disorders such as depression and chronic pain.

Introduction
Just over fifty years ago, psychologists James Olds and Peter Milner, working at McGill
University in Canada, carried out their pioneering experiments which discovered that rats
would repeatedly press levers to receive tiny jolts of current injected through electrodes
implanted deep within their brains (Olds and Milner, 1954). Especially when this brain
stimulation was targeted at certain areas of the brain in the region of the septum and nucleus
accumbens, the rats would repeatedly press the lever -- even up to 2000 times per hour
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(Olds, 1956).

These powerful findings seemed to suggest that Olds and Milner had discovered the pleasure
center in the brain. Research in the next two decades established that dopamine is one of the
main chemicals aiding neural signaling in these regions, and for many years dopamine was
suggested to be the brain’s “pleasure chemical.” The results seemed to promise an easy fix
to the unhappiness and suffering which is the traveling companion of far too many people.
They certainly emboldened writers to envisage brave new worlds where drugs and electrical
stimulation could induce bliss for the masses.

But is the high road to happiness really that simple? Subsequent human experiments suggest
otherwise. Around the same time in the 1950s and 1960s, American psychiatrist Robert
Heath at Tulane University took it upon himself to further these findings in some ethically
questionable experiments on mentally ill human patients (Baumeister, 2000). Infamously, in

Corresponding authors: Morten L. Kringelbach, D.Phil. (morten.kringelbach@psych.ox.ac.uk) and Kent C. Berridge, Ph.D.
(berridge@umich.edu).
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one case he even implanted electrodes to try to cure homosexuality (Heath, 1972). This line
of research was eventually stopped. Most substantively, however, the pleasure electrodes
may never have lived up to their name. Although the researchers also found compulsive
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lever pressing in some patients, it was never clear from these patients’ subjective reports that
the electrodes did indeed cause real pleasure. Some researchers today suggest that the
electrodes never caused intense pleasure or ‘liking’ after all, but only a form of ‘wanting’ or
motivation to obtain the stimulation (see discussion in Green et al., 2010; Smith et al.,
2010).

Pleasure and happiness are linked, however, but in much more complex ways than simple
pleasure electrodes would suggest, even if such electrodes exist. In this review we map out
some of the intricate links between them to show how they are at the heart of affective
neuroscience and the psychology of well-being. We will synthesize the results of last fifty
years of careful study of reward and affective processing in the brain. Our main contention
is that a better understanding of the pleasures of the brain may offer a more general insight
into happiness, into how brains work to produce it in daily life for the fortunate, how brains
fail in the less fortunate, and hopefully into better ways to enhance the quality of life.

A Science of Pleasure
The scientific study of pleasure and affect was pioneered by the ideas of Charles Darwin,
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who examined the evolution of emotions and affective expressions, and suggested that
affects are adaptive responses to environmental situations. Prominent affective reactions
such as pleasure ‘liking’ and displeasure reactions can be found in the behavior and brains of
all mammals (Steiner et al., 2001), and likely have important evolutionary functions
(Kringelbach, 2009). Both positive affect and negative affect have been proposed to have
adaptive functions (Nesse, 2004) and it is clear that the neural mechanisms for generating
affective reactions are present and similar in most mammalian brains, and as such appear to
have been long ago evolutionarily selected for and conserved across species from humans to
rodents (Kringelbach, 2010).

The progress in affective neuroscience in recent years has been made possible by identifying
objective aspects of pleasure-elicited reactions and triangulating toward underlying brain
substrates. This scientific strategy divides the concept of affect into two parts: the affective
state, which has objective aspects in behavioral, physiological, and neural reactions; and
conscious affective feelings, seen as the subjective experience of emotion (Kringelbach,
2004a). This definition allows conscious feelings to play a central role in hedonic
experiences, but holds that the affective essence of a pleasure reaction is not limited to this
conscious feeling. It means that objective affective state can be measured in other animals,
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regardless of the availability or accuracy of corresponding subjective reports, and as such is

especially tractable to neuroscience investigations that involve brain manipulations.

The available evidence suggests that brain mechanisms involved in fundamental pleasures
(food and sexual pleasures) overlap with those for higher-order pleasures (for example,
monetary, artistic, musical, altruistic, and transcendent pleasures) (Kringelbach, 2010).

It is an important hedonic principle that the rewarding properties for all pleasures are likely
to be generated by hedonic brain circuits that are distinct from the mediation of other
features of the same events (e.g., sensory, cognitive) (Kringelbach, 2005). Thus pleasure is
never merely a sensation or a thought, but is instead an additional hedonic gloss generated
by the brain via dedicated systems (Frijda, 2010).

All pleasures from sensory pleasures and drugs of abuse to monetary, aesthetic, and musical
delights would seem to involve the same fundamental hedonic brain systems. Pleasures

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important to happiness, such as socializing with friends, and related traits of positive
hedonic mood, are thus all likely to draw upon the same neurobiological roots that evolved
for sensory pleasures. The neural overlap may offer a way to generalize from fundamental
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pleasures that are best understood and so infer larger hedonic brain principles likely to
contribute to happiness.

The Neuroanatomy of Pleasure

Pleasure is a complex psychological concept with many different sub-components which
include ‘liking,’ ‘wanting,’ and ‘learning’ components (Berridge and Kringelbach, 2008;
Smith et al., 2010). Each component has both conscious and non-conscious elements that
can be studied in humans -- and at least the latter can also be probed in other animals (Figure
1).

Hedonic hotspots
The brain has an extensive distribution of reward-related circuitry with some hedonic
mechanisms found deep in the brain (nucleus accumbens, ventral pallidum, brainstem) and
other candidates are in the cortex (orbitofrontal, cingulate, medial prefrontal, and insular
cortices) (Figure 2). Pleasure coding brain networks are widespread and provide evidence
for highly distributed brain coding of hedonic states. Yet, pleasure causation, which can be
detected as increases in ‘liking’ reactions consequent to brain manipulation, has so far been
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found for only a few hedonic hotspots in the subcortical structures. Each hedonic hotspot is
merely a cubic-millimeter or so in volume in the rodent brain (and should be a cubic-
centimeter or so in humans, if proportional to whole brain volume). Hotspots are capable of
generating enhancements of ‘liking’ reactions to a sensory pleasure such as sweetness, when
stimulated with opioid, endocannabinoid, or other neurochemical modulators (Smith et al.,
2010).

Hotspots have been found in nucleus accumbens shell and ventral pallidum, and possibly
other forebrain and limbic cortical regions, and also in deep brainstem regions including the
parabrachial nucleus in the pons (Figure 2D). The pleasure-generating capacity of these
hotspots has been revealed in part by studies in which microinjections of drugs stimulated
neurochemical receptors on neurons within a hotspot, and caused a doubling or tripling of
the number of hedonic ‘liking’ reactions normally elicited by a pleasant sucrose taste.
Analogous to scattered islands that form a single archipelago, hedonic hotspots are
anatomically distributed but interact to form a functional integrated circuit. The circuit obeys
control rules that are largely hierarchical and organized into brain levels. Top levels function
together as a cooperative heterarchy, so that, for example, multiple unanimous ‘votes’ in
favor from simultaneously-participating hotspots in the nucleus accumbens and ventral
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pallidum are required for opioid stimulation in either forebrain site to enhance ‘liking’ above
normal.

In addition, as mentioned above, pleasure is translated into motivational processes in part by

activating a second component of reward termed ‘wanting’ or incentive salience, which
makes stimuli attractive when attributed to them by mesolimbic brain systems (Berridge and
Robinson, 2003). Incentive salience depends in particular on mesolimbic dopamine
neurotransmission (though other neurotransmitters and structures also are involved).

Importantly, incentive salience is not hedonic impact or pleasure ‘liking’ (Berridge, 2007).
This is why an individual can ‘want’ a reward without necessarily ‘liking’ the same reward.
Irrational ‘wanting’ without liking can occur especially in addiction via incentive-
sensitization of the mesolimbic dopamine system and connected structures. At extreme, the
addict may come to ‘want’ what is neither ‘liked’ nor expected to be liked, a dissociation

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possible because ‘wanting’ mechanisms are largely subcortical and separable from
cortically-mediated declarative expectation and conscious planning. This is a reason why
addicts may compulsively ‘want’ to take drugs even if, at a more cognitive and conscious
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level, they do not want to do so. That is surely a recipe for great unhappiness (Figure 2,
bottom right).

Cortical pleasure
Hedonic evaluation of pleasure valence is separable from precursor operations such as
sensory computations, suggesting existence of a hedonic cortex proper (Figure 2). Hedonic
cortex involves regions such as the orbitofrontal, insula, medial prefrontal, and cingulate
cortices, which, shown by a wealth of human neuroimaging studies, code for hedonic
evaluations (including anticipation, appraisal, experience, and memory of pleasurable
stimuli) and have close anatomical links to subcortical hedonic hotspots. It is important,
however, to again make a distinction between brain activity coding and causing pleasure.
Neural coding is inferred in practice by measuring brain activity correlated to a pleasant
stimulus, using human neuroimaging techniques, or electrophysiological or neurochemical
activation measures in animals (Aldridge and Berridge, 2010). Causation is generally
inferred on the basis of a change in pleasure as a consequence of a brain manipulation such
as a lesion or stimulation. Coding and causation often go together for the same substrate, but
they may diverge so that coding occurs alone.
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In humans, pleasure encoding may reach an apex of cortical localization in a subregion that
is mid-anterior and roughly mid-lateral within the orbitofrontal cortex of the prefrontal lobe,
where neuroimaging activity correlates strongly to subjective pleasantness ratings of food
varieties - and to other pleasures such as sexual orgasms, drugs, chocolate, and music. Most
importantly, activity in this special mid-anterior zone of orbitofrontal cortex tracks changes
in subjective pleasure, such as a decline in palatability when the reward value of one food
was reduced by eating it to satiety (while remaining high to another food). The mid-anterior
subregion of orbitofrontal cortex is thus a prime candidate for the coding of subjective
experience of pleasure (Kringelbach, 2005).

Another potential coding site for positive hedonics in orbitofrontal cortex is along its medial
edge that has activity related to the positive and negative valence of affective events
(Kringelbach and Rolls, 2004), contrasted to lateral portions that have been suggested to
code unpleasant events (although lateral activity may reflect a signal to escape the situation,
rather than displeasure per se) (Kringelbach, 2004b; O’Doherty et al., 2001). This medial-
lateral hedonic gradient interacts with an abstraction-concreteness gradient in the posterior-
anterior dimension, so that more complex or abstract reinforcers (such as monetary gain and
loss) are represented more anteriorly in the orbitofrontal cortex than less complex sensory
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rewards (such as taste). The medial region that codes pleasant sensations does not, however,
appear to change its activity with reinforcer devaluation, and so may not reflect the full
dynamics of pleasure.

Still other cortical regions have been implicated by some studies in coding for pleasant
stimuli, including parts of the mid-insular cortex that is buried deep within the lateral surface
of the brain as well as parts of the anterior cingulate cortices on the medial surface of the
cortex (Kringelbach, 2005). As yet, however, pleasure coding is not as clear for those
regions as for the orbitofrontal cortex, and it remains uncertain whether insular or anterior
cingulate cortices specifically code pleasure or only emotion more generally.

It remains still unknown, however, if even the mid-anterior pleasure-coding site of

orbitofrontal cortex or medial orbitofrontal cortex or any other cortical region actually
causes a positive pleasure state. Clearly, damage to orbitofrontal cortex does impair

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pleasure-related decisions, including choices and context-related cognitions in humans,

monkeys, and rats (Anderson et al., 1999; Nauta, 1971). But some caution regarding
whether cortex generates positive affect states per se is indicated by the consideration that
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patients with lesions to the orbitofrontal cortex do still react normally to many pleasures,
although sometimes showing inappropriate emotions. Hedonic capacity after prefrontal
damage has not, however, yet been studied in careful enough detail to draw firm conclusions
about cortical causation (e.g., using selective satiation paradigms), and it would be useful to
have more information on the role of orbitofrontal cortex, insular cortex, and cingulate
cortex in generating and modulating hedonic states.

Pleasure causation has been so far rather difficult to assess in humans given the limits of
information from lesion studies, and the correlative nature of neuroimaging studies. A
promising tool, however, is deep brain stimulation (DBS) which is a versatile and reversible
technique that directly alters brain activity in a brain target and where the ensuing whole-
brain activity can be measured with MEG (Kringelbach et al., 2007). Pertinent to a view of
happiness as freedom from distress, at least pain relief can be obtained from DBS of
periaqueductal grey in the brainstem in humans, where specific neural signatures of pain
have been found (Green et al., 2009), and where the pain relief is associated with activity in
the mid-anterior orbitofrontal cortex, perhaps involving endogenous opioid release.
Similarly, DBS may alleviate some unpleasant symptoms of depression, though without
actually producing positive affect.
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Linking Pleasure and Happiness

Pleasure can thus be seen to drive life, as most animals know it by the rewards associated
with fulfilling ancient evolutionary imperatives of survival and procreation. Humans of
course are able to consciously experience these pleasures and, perhaps uniquely, even
contemplate the elusive prospect of happiness.

The advanced human ability to consciously predict and anticipate the outcome of choices
and actions confers our species with an evolutionary advantage, but human conscious
planning is a double-edged sword as John Steinbeck pointed out as he wrote of “the tragic
miracle of consciousness” and how our “species is not set, has not jelled, but is still in a state
of becoming” (Steinbeck and Ricketts, 1941). While consciousness allows us to experience
pleasures, desires, and perhaps even happiness, this is always accompanied by the certainty
of the end; yet most people remain optimistic in the face of adversity.

Happiness is, however, a slippery concept (Gilbert, 2006; Bloom, 2010). One way to
approach it is to follow the insight of Aristotle that happiness can usefully thought of as
consisting of two fundamental aspects: hedonia (pleasure) and eudaimonia (a life well-
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lived). In contemporary psychology these aspects are usually referred to as pleasure and
meaning, and positive psychologists have recently proposed to add a third meaning-related
component of engagement involving feelings of commitment and participation in life
(Seligman et al., 2005).

Using these definitions scientists have made substantial progress in defining and measuring
happiness in the form of self-reports of subjective well-being, in identifying its distribution
across people in the real world, and in identifying how well-being is influenced by various
life factors ranging from income to other people (Kahneman, 1999). This research shows
that while there is clearly a sharp conceptual distinction between pleasure versus
engagement-meaning components, hedonic and eudaimonic aspects empirically cohere
together in happy people (Diener et al., 2006; Kahneman, 1999; Seligman et al., 2005).

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Surveys of happiness provide interesting indicators of mental well-being in societies, but

offer little evidence of the underlying neurobiology of happiness. Supporting a hedonic
approach to that question, it has been suggested that the best measure of subjective well-
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being may be simply to ask people how they hedonically feel right now -- again and again --
so as to track their hedonic accumulation across daily life (Kahneman, 1999). Such repeated
self-reports of hedonic states could also be used to identify more stable neurobiological
hedonic brain traits that dispose particular individuals toward happiness. Further, a hedonic
approach might even offer a toehold into identifying eudaimonic brain signatures of
happiness, due to the empirical convergence between the two categories, even if pleasant
mood is only half the happiness story (Kringelbach and Berridge, 2009).

We have previously suggested that one possible toehold linking pleasure and happiness
might be found in the close links between sensory pleasure networks and the brain’s default
network (Kringelbach and Berridge, 2009) (Figure 3). We have proposed that the
eudaimonic happiness may be linked to potential interactions of hedonic brain circuits with
circuits that assess meaningful relationships of self to social others (Lou et al., 1999),
internal modes of cognition (Buckner et al., 2008), and perhaps even states of consciousness
(Laureys et al., 2004). The default network might deserve further consideration for a role in
connecting eudaimonic and hedonic happiness. At least, key regions of the frontal default
network overlap with the hedonic network, such as the anterior cingulate and orbitofrontal
cortices, and have a relatively high density of opiate receptors. Similarly, activity changes in
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the frontal default network, such as in the subgenual cingulate and orbitofrontal cortices,
correlate to pathological changes in subjective hedonic experience, such as in depressed
patients (Drevets et al., 1997).

Pathological self-representations by the frontal default network could also contribute in

unfortunate individuals to hedonic distortions of happiness that involve eudaimonic
dissatisfaction, such as in cognitive rumination of depression. Conversely, mindfulness-
based cognitive therapy for depression, which aims to disengage from dysphoria-activated
depressogenic thinking might conceivably recruit default network circuitry to help mediate
improvement in happiness via a linkage of eudaimonic to hedonic circuitry.

Conclusions
At first glance, pleasure electrodes once seemed to provide the prospect of happiness at the
flick of a switch but careful scientific experimentation has shown that such electrodes are
unlikely to truly cause pleasure, and are instead likely linked most closely to the
psychological processes of ‘wanting’ -- with very little ‘liking’ involved (Berridge and
Kringelbach, 2008).
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Both pleasure and happiness are much more complex psychological states than the unitary
words imply, with multiple sub-components within each; some of which are amenable to
scientific investigation even now. In this article, we have shown the progress in building a
science of pleasure and we have identified some of the mechanisms and regions important in
the brain’s hedonic networks that generate basic pleasures. We have also speculated on
potential interaction of hedonics with eudaimonic networks that may be important
contributors to happiness. Yet, it is important to note that we have still not made substantial
progress towards understanding the functional neuroanatomy of happiness.

While it remains unclear how pleasure and happiness are exactly linked, it may be safe to
say at least that the pathological lack of pleasure, in anhedonia or dysphoria, amounts to a
formidable obstacle to happiness. Exciting new insights have been gained while studying
sensory pleasures, but many further challenges remain such as to understand how the brain

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networks underlying fundamental pleasure relate to higher pleasures such as music, dance,
play, and flow to contribute to happiness.
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Further, in social animals like humans, it is worth noting that cultural interactions with
conspecifics are fundamental and central to enhancing the other pleasures. Humans are
intensely social, and data indicate that one of the most important factors for happiness is
relationships with other people. Social pleasures may still include vital sensory features such
as visual faces, touch features of grooming and caress, as well as in humans more abstract
and cognitive features of social reward and relationship evaluation. These may be especially
important triggers for the brain’s hedonic networks in human beings.

In particular, adult pair bonds and attachment bonds between parents and infants are likely
to be extremely important for the survival of the species (Kringelbach et al., 2008). The
breakdown of these bonds is all too common and can lead to great unhappiness. And even
bond formation can potentially disrupt happiness, such as in transient parental depression
after birth of an infant - in over 10% of mothers and approximately 3% of fathers (Cooper
and Murray, 1998). Progress in understanding the hedonics of social bonds could be useful
in understanding happiness, and it will be important to map the developmental changes that
occur over a lifespan. Fortunately, social neuroscience is beginning to unravel some of the
complex dynamics of human social interactions and their relation to brain activations
(Parsons et al., 2010).
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Many future challenges remain before we will understand the functional neuroanatomy of
happiness. We have previously proposed that hedonic happiness could be akin to ‘liking’
without ‘wanting;’ as a state of pleasure without disruptive desires -- a state of contentment
(Kringelbach, 2009). Yet, alternatively happiness in daily life may rely on matching a proper
balance ‘wanting’ and ‘liking’ to help facilitate engagement with the world. If the balance
tips the wrong way, happiness becomes impossible. As an example too much ‘wanting’ can
readily spiral into maladaptive patterns such as addiction, and is a certain recipe to great
unhappiness. And of course, the eudaimonic components of meaning and engagement are
crucial to happiness for human beings. Careful scientific experimentation will create a better
scientific understanding of pleasure and happiness that may someday allow clinicians to
make targeted interventions that will help to shift more among us into a better situation to
enjoy daily events, to find life meaningful and worth living -- and perhaps even to achieve a
degree of bliss.

Acknowledgments
Our research has been supported by grants from the TrygFonden Charitable Foundation to MLK and from the
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NIMH and NIDA to KCB.

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Figure 1.
A scientific program for the study of pleasure. Pleasure is a complex psychological concept
with at least three major subcomponents of motivation or wanting (white), pleasure liking or
affect (light blue), and learning (blue). Each of these contains explicit (top rows, light
yellow) and implicit (bottom rows, yellow) psychological components (second column) that
constantly interact and require careful scientific experimentation to tease apart. Explicit
processes are consciously experienced (e.g., explicit pleasure and happiness, desire, or
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expectation), whereas implicit psychological processes are potentially unconscious in the

sense that they can operate at a level not always directly accessible to conscious experience
(implicit incentive salience, habits, and ‘liking’ reactions), and must be further translated by
other mechanisms into subjective feelings. Measurements or behavioral procedures that are
especially sensitive markers of the each of the processes are listed (third column). Examples
of some of the brain regions and neurotransmitters are listed (fourth column), as well as
specific examples of measurements (fifth column), such as an example of how highest
subjective life satisfaction does not lead to the highest salaries (top) (Haisken-De New and
Frick, 2005). Another example shows the incentive-sensitization model of addiction and
how ‘wanting’ to take drugs may grow over time independently of ‘liking’ and ‘learning’
drug pleasure as an individual becomes an addict (bottom) (Robinson and Berridge, 1993).
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Figure 2.
Hedonic brain circuitry in humans and other animals. Pleasure-elicited reactions allow us to
investigate the brain regions involved in pleasure in rodents and humans. (a) Facial ‘liking’
and ‘disliking’ expressions elicited by sweet and bitter taste are similar in rodents and
human infants. (b, d) Pleasure causation has been identified in rodents as arising from
interlinked subcortical hedonic hotspots, such as in nucleus accumbens and ventral pallidum,
where neural activation may increase ‘liking’ expressions to sweetness. Similar pleasure
coding and incentive salience networks have also been identified in humans. (c) The so-
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called ‘pleasure’ electrodes in rodents and humans are unlikely to have elicited true pleasure
but perhaps only incentive salience or ‘wanting.’ (d) The cortical localization of pleasure
coding may reach an apex in various regions of the orbitofrontal cortex, which differentiate
subjective pleasantness from valence processing of aspects of the same stimulus, such as a
pleasant food.
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Figure 3.
A hypothesis of how pleasure and happiness are linked. Default networks are fundamental to
human brain function and have been linked to self awareness, remembering the past and
prospecting the future (a-c). It is clear that these networks are partly overlapping the
pleasure networks. We have hypothesized that happiness might include a role for the default
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network, or for related neural circuits that contribute to computing relations between self
and others, in evaluating eudaimonic meaning and interacting with hedonic circuits of
positive affect. Some examples show (d) key regions of the default network such as the
anterior cingulate and orbitofrontal cortices that have a high density of opiate receptors, (e)
have been linked to depression, and (f) its surgical treatment. (g) Subregional localization of
function may be indicated by connectivity analyses of cingulate cortex and related
structures, (h) important in pleasure-related monitoring, learning, and memory, (i) as well as
self-knowledge, person perception, and other cognitive functions. (j) The default network
may change over early life in children and pre-term babies, (k) in pathological states
including depression and vegetative states, (l) and after lesions to its medial orbitofrontal
and subgenual cingulate cortices that disrupt reality monitoring and create spontaneous
confabulations.
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