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1 chapter 

2 the neurobiology 3 of and 4

5 morten l. kringelbach and kent c. berridge

6 Introduction

7 Happiness is an elusive state, diffi cult to defi ne, and therefore challenging to measure— 8 partly due to its clearly subjective, and perhaps uniquely , . But how can one 9 get a scientifi c handle on such a slippery concept? 10 Since , happiness has been thought of as consisting of at least two aspects: 11 hedonia (pleasure) and (a life well-lived) (Waterman 1993 ). In contemporary 12 psychology these aspects are usually referred to as pleasure and meaning, and scientists have 13 recently proposed to add a third distinct component of engagement related to of 14 commitment and participation in life (Seligman et al. 2 0 0 5 ) . 15 Using these defi nitions scientists have made substantial progress in defi ning and measur- 16 ing happiness in the form of self-reports of subjective well-being (Kahneman 1999 ; Ryan 17 and Deci 2001 ; Diener et al. 2003 ; Seligman et al. 2 0 0 5 ) . Th is research shows that while there 18 is clearly a sharp conceptual distinction between pleasure versus engagement-meaning 19 components, hedonic and eudaimonic aspects empirically cohere together in happy people. 20 For example, in happiness surveys over 80 of people rate their overall eudaimonic life 21 satisfaction as “pretty to very happy”, and comparably, 80  also rate their current hedonic 22 mood as positive (e.g. positive 6–7 on a 10-point scale where 5 is hedonically neutral) 23 (Kesebir and Diener 2008 ). A lucky few may even live consistently around a hedonic point 24 of 8 — although excessively higher hedonic scores may actually impede attainment of life 25 success, as measured by riches, education, or political participation (Oishi et al. 2 0 0 7 ) . 26 While these surveys are interesting indications of mental well-being, they off er little 27 evidence of the underlying neurobiology of happiness. In this review we will therefore focus 28 on the substantial progress in understanding the psychology and neurobiology of sensory 29 pleasure that has been made over the last decade (Berridge and Kringelbach 2008 ; 30 Kringelbach and Berridge 2010 ).

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 morten l. kringelbach and kent c. berridge

1 Th ese advances make the hedonic side of happiness most tractable to a scientifi c approach 2 to the neural underpinnings of happiness. Supporting a hedonic approach, it has been sug- 3 gested that the best measure of subjective well-being may be simply to ask people how they 4 hedonically feel right now — again and again — so as to track their hedonic accumulation 5 across daily life (Kahneman 2000 ; Diener et al. 2003 ; Gilbert and Wilson 2007 ). Th ese 6 repeated self-reports of hedonic states could also be used to identify more stable neurobio- 7 logical hedonic brain traits that dispose particular individuals toward happiness. Further, a 8 hedonic approach might even off er a toehold into identifying eudaimonic brain signatures 9 of happiness, due to the empirical convergence between the two categories, even if pleasant 10 mood is only half the happiness story (Kringelbach and Berridge 2009 ). 11 It is important to note that our focus on the hedonia component of happiness should not 12 be confused with , which is the pursuit of pleasure for pleasure’s own sake, and 13 more akin to the features we describe later. Also, to focus on hedonics does not 14 deny that some ascetics may have found bliss through painful self-sacrifi ce, but simply 15 refl ects that positive hedonic tone is indispensable to most people seeking happiness.

16 A of pleasure

17 Th e link between pleasure and happiness has a long history in psychology. It was stressed in 18 the early writings of (Freud and Riviere 1930 ), when he posited that people 19 “strive aft er happiness; they want to become happy and to remain so. Th is endeavor has 20 two sides, a positive and a negative aim. It aims, on the one hand, at an absence of and 21 displeasure, and, on the other, at the experiencing of strong feelings of pleasure” (Freud and 22 Riviere 1930 , p. 76). Emphasizing a positive balance of aff ect to be happy implies that studies 23 of hedonic brain circuits can advance the of both pleasure and happiness. 24 A related but slightly diff erent view is that happiness depends most chiefl y on eliminating 25 negative “pain and displeasure” to free an individual to pursue engagement and meaning. 26 Positive pleasure by this view is somewhat superfl uous. Th is view may characterize the 20 th 27 century medical and clinical emphasis on alleviating negative psychopathology and strongly 28 distressing . It fi ts also with ’s early quip that “Happiness, I have lately 29 discovered, is no positive , but a negative condition of freedom from a number of 30 restrictive sensations of which our organism usually seems the seat. When they are wiped 31 out, the clearness and cleanness of the contrast is happiness. Th is is why anaesthetics make 32 us so happy. But don’t you take to drink on that account.” (James 1920 , vol 2, p. 158). 33 Focusing on eliminating negative distress seems to leave positive pleasure outside the 34 boundary of happiness, perhaps as an extra bonus or even an irrelevancy for ordinary pur- 35 suit. In practice, many mixtures of positive aff ect and negative aff ect may occur in individu- 36 als (Ryff et al. 2006 ) and cultures may vary in the importance of positive versus negative 37 aff ect for happiness. For example, positive emotions are linked most strongly to ratings of 38 life satisfaction overall in nations that stress self-expression, but alleviation of negative emo- 39 tions may become relatively more important in nations that (Kuppens 40 et al. 2008 ). 41 By either view, psychology seems to be moving away from the stoic notion that aff ect 42 states such as pleasure are simply irrelevant to happiness. Th e growing evidence for the

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the neurobiology of pleasure and happiness 

1 importance of aff ect in psychology and neuroscience shows that a scientifi c account will 2 have to involve hedonic and/or displeasures. To move towards a neuroscience of 3 happiness, a neurobiological understanding is required of how positive and negative aff ect 4 are balanced in the brain. 5 Given the potential contributions of hedonics to happiness, we now survey develop- 6 ments in understanding brain mechanisms of pleasure (Berridge and Kringelbach 2008 ; 7 Leknes and Tracey 2008 ). Th e scientifi c study of pleasure and aff ect was foreshadowed by 8 the pioneering ideas of Charles Darwin, who examined the evolution of emotions and 9 aff ective expressions, and suggested that these are adaptive responses to environmental sit- 10 uations. In that vein, pleasure “liking” and displeasure reactions are prominent aff ective 11 reactions in the behavior and brains of all mammals (Steiner et al. 2001 ), and likely had 12 important evolutionary functions (Kringelbach 2009 ). Neural mechanisms for generating 13 aff ective reactions are present and similar in most mammalian brains, and thus appear 14 to have been selected for and conserved across species (Kringelbach 2010 ). Indeed, both 15 positive aff ect and negative aff ect are recognized today as having adaptive functions (Nesse 16 2004 ), and positive aff ect in particular has consequences in daily life for planning and 17 building cognitive and emotional resources (Fredrickson et al. 2008 ; Dickinson and 18 Balleine 2010 ). 19 Such functional perspectives suggest that aff ective reactions may have objective features 20 beyond subjective ones (Kringelbach 2004a ). Progress in aff ective neuroscience has been 21 made recently by identifying objective aspects of pleasure reactions and triangulating 22 toward underlying brain substrates. Th is scientifi c strategy divides the concept of aff ect into 23 two parts: the aff ective state , which has objective aspects in behavioral, physiological, and 24 neural reactions; and conscious aff ective feelings , seen as the subjective experience of 25 (Kringelbach 2004a ). Note that such a defi nition allows conscious feelings to play a central 26 role in hedonic experiences, but holds that the aff ective essence of a pleasure reaction is 27 more than a conscious feeling. 28 Evidence so far available suggests that brain mechanisms involved in fundamental plea- 29 sures (food and sexual pleasures) overlap with those for higher-order pleasures (for example, 30 monetary, artistic, musical, altruistic, and transcendent pleasures) (Small et al. 2 0 0 1 ; 31 Kahneman et al. 2004 ; Kringelbach 2005 ; Peciña et al. 2006 ; Gottfried 2010 ; Kringelbach 32 2010 ; Kringelbach et al. 2010 ; Veldhuizen et al. 2010 ). 33 From sensory pleasures and drugs of abuse (Robinson and Berridge 2003 ) to monetary, 34 aesthetic, and musical delights, all pleasures seem to involve the same hedonic brain sys- 35 tems, even when linked to and memory (Skov 2010 ; Vuust and Kringelbach 36 2010 ). Pleasures important to happiness, such as socializing with friends (Kahneman 1999 ; 37 Ryan and Deci 2001 ; Diener et al. 2 0 0 3 ; K a h n e m a n et al. 2004 ; Seligman et al. 2 0 0 5 ) , a n d 38 related traits of positive hedonic mood are thus all likely to draw upon the same neurobio- 39 logical roots that evolved for sensory pleasures. Th e neural overlap may off er a way to gener- 40 alize from fundamental pleasures that are best understood and so infer larger hedonic brain 41 principles likely to contribute to happiness. 42 We note the rewarding properties for all pleasures are likely to be generated by hedonic 43 brain circuits that are distinct from the mediation of other features of the same events (e.g. 44 sensory, cognitive) (Kringelbach 2005 ). Th us pleasure is never merely a sensation or a 45 thought (Frijda 2010 ), but is instead an additional hedonic gloss generated by the brain via 46 dedicated systems.

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 morten l. kringelbach and kent c. berridge

1 The of pleasure

2 How does positive aff ect arise? Aff ective neuroscience research on sensory pleasure has 3 revealed many networks of brain regions and activated by pleasant events 4 and states (Figures 2.1 and 2.2 ). Identifi cation of hedonic substrates has been advanced by 5 recognizing that pleasure or “liking” is but one component in the larger composite psycho- 6 logical process of reward, which also involves “wanting” and “” components (Smith 7 et al. 2010 ). Each component also has conscious and non-conscious elements that can be 8 studied in — and at least the latter can also be probed in other animals.

9 Hedonic hotspots

10 Despite having an extensive distribution of reward-related circuitry, the brain appears rather 11 frugal in “liking” mechanisms that cause pleasure reactions. As shown in later paragraphs, 12 some hedonic mechanisms are found deep in the brain (, ventral pal- 13 lidum, brainstem) and other candidates are in the cortex (orbitofrontal, cingulate, medial 14 prefrontal, and insular cortices) (Berridge 1996 ; Cardinal et al. 2002 ; Kringelbach et al. 2 0 0 3 ; 15 Kringelbach and Rolls 2004 ; Everitt and Robbins 2005 ; Amodio and Frith 2006 ; Kringelbach 16 2010 ; Watson et al. 2010 ). Pleasure-activated brain networks are widespread, but compelling 17 evidence for pleasure causation (detected as increases in “liking” reactions consequent to 18 brain manipulation) has so far been found for only a few hedonic hotspots in the subcortical 19 structures. Each hotspot is merely a cubic millimeter or so in volume in the rodent brain 20 (and should be a cubic centimeter or so in humans, if proportional to whole brain volume). 21 Hotspots are capable of generating enhancements of “liking” reactions to a sensory pleasure 22 such as , when stimulated with , endocannabinoid, or other neurochemical 23 modulators (Smith et al. 2010 ). 24 Hotspots exist in nucleus accumbens shell and ventral pallidum, and possibly other fore- 25 brain and limbic cortical regions, and also in deep brainstem regions including the parabra- 26 chial nucleus in the pons (Figure 2.2d ) (Peciña et al. 2 0 0 6 ) . Th e pleasure-generating capacity 27 of these hotspots has been revealed in part by studies in which microinjections of drugs 28 stimulated neurochemical receptors on neurons within a hotspot, and caused a doubling or 29 tripling of the number of hedonic “liking” reactions normally elicited by a pleasant sucrose 30 taste (Smith et al. 2010 ). Analogous to scattered islands that form a single archipelago, hedo- 31 nic hotspots are anatomically distributed but interact to form a functional integrated circuit. 32 Th e circuit obeys control rules that are largely hierarchical and organized into brain levels. 33 Top levels function together as a cooperative heterarchy, so that, for example, multiple 34 unanimous “votes” in favor from simultaneously-participating hotspots in the nucleus 35 accumbens and ventral pallidum are required for opioid in either forebrain site 36 to enhance “liking” above normal (Smith and Berridge 2007 ). 37 In addition, as mentioned earlier, pleasure is translated into motivational processes in 38 part by activating a second component of reward termed “wanting” or incentive salience , 39 which makes stimuli attractive when attributed to them by mesolimbic brain systems 40 (Berridge and Robinson 2003 ). Incentive salience depends in particular on mesolimbic

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the neurobiology of pleasure and happiness  c rst

Explicit Implicit Berridge ll as specifi ll as tantly interact interact tantly ach of the pro- of ach e at a level not not a level e at rther translated by

Each gray). ect (light Time

“Liking” “wanting” “Learning” Life satisfaction (1984–1987) cortex; NAc, nucleus accumbens; accumbens; nucleus NAc, cortex; Examples of measurements 0

5,000

25,000 20,000 15,000 10,000 Salary (2000-2003) Salary prefrontal OFC, ACC, insular OFC, ACC, insular , OFC, ACC, mPFC, insular Ach, dopamine, NAc, VTA, dopamine PAG, NAc shell, VP, opioids, cannabinoids Amygdala, hippocampus Ach, dopamine oral procedures that are especially sensitive markers of the e of markers especially sensitive are that procedures oral Subjective ratings of cognitive goals Subjective ratings of pleasure Rational inference verbal explanation Conditioned approach, PIT autoshaping, cued relapse Facial affective expressions Facial affective human pleasure-elicited reactions Pavlovian conditioned response response reinforcement instr. idum;VTA, ventral tegmental area. area. tegmental ventral idum;VTA, Wanting cognitive incentives Liking conscious pleasure Learning cognitive processing “Wanting” incentive salience “Liking” hedonic impact “Learning” associative learning h column), such as an example of how highest subjective life satisfaction does not lead to the highest salaries (top) leadthe doeshighest salaries to not (top) satisfaction life highest subjective how of example as an such h column), Psychological componentsPsychological Measurements Examples of brain circuitry ft c experimentation to tease apart. Explicit processes are consciously experienced (e.g. explicit pleasure and happiness, happiness, and pleasure experienced explicit (e.g. consciously are processes tease Explicit to apart. c experimentation Major categories Motivation Pleasure Learning Measuring reward and hedonia. Reward and pleasure are multifaceted psychological concepts. Major processes within reward (fi reward within processes Major concepts. psychological multifaceted are pleasure and Reward hedonia. and reward Measuring examples of measurements (fi measurements of examples column) consist of motivation or wanting (white), learning (gray), and — most relevant to happiness — pleasure liking or aff or liking pleasure — happiness to most relevant — and (gray), learning (white), wanting or motivation of consist column) fig. 2.1 of these contains explicit (top three rows) and implicit (bottom three rows) psychological components (second column) that cons that column) (second components psychological rows) three (bottom implicit and rows) three (top explicit these contains of and require careful scientifi careful require and they operat in the can that unconscious potentially are processes psychological implicit expectation), whereas or desire, be fu must and reactions), “liking” and salience, habits incentive experience (implicit conscious to accessible directly always take to drugs may “wanting” how modeladdiction and of the incentive-sensitization shows example 2005). Another Frick and New (Haisken-De and (Robinson addict becomes(bottom) an individual as an drug pleasure “learning” and “liking” of time independently over mPFC, medial cortex; grow orbitofrontal OFC, ; Ach, cortex; cingulate anterior ). ACC, 1993 pall ventral gray; VP, periaqueductal PAG, other mechanisms into subjective feelings. Measurements or behavi or Measurements feelings. subjective into mechanisms other cesses are listed (third column). Examples of some of the brain regions and neurotransmitters are listed (fourth column), as we (fourth column), listed are neurotransmitters and regions the brain of some of Examples column). (third listed cesses are

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 morten l. kringelbach and kent c. berridge tive saliencetive “Liking” decrease Hedonic coldspot s and human infants. infants. human s and “Disliking” decrease Medial OFC Mid-anterior OFC “Liking” increase Pleasure causation and coding increase hotspot “Wanting” D Hedonic hotspot 1mm 10mm ng of aspects the same stimulus, such as a pleasant as a pleasant such aspects of ng the stimulus, same cal hedonic hotspots, such as in nucleus accumbens as in nucleus such hotspots, cal hedonic Pleasure electrodes C ” expressions to sweetness. Similar pleasure coding and incen and coding pleasure Similar sweetness. to ” expressions gure shows the brain regions for causing and coding fundamental pleasure pleasure coding fundamental and causing for regions the brain shows gure ons elicited by sweet and bitter taste are similar in rodent similar are taste bitter and sweet by elicited ons VTA Cingulate cortex Hypothalamus PAG Nucleus accumbens pallidum Ventral Amygdala regions in various apex an reach coding may pleasure of e cortical localization Liking and wanting regions fi e schematic true elicited have to unlikely are humans and electrodes in rodents e so-called “pleasure” Hedonic brain circuits ed in rodents as arising from interlinked subcorti interlinked from as arising ed in rodents erentiate subjective pleasantness from valence processi valence from pleasantness subjective erentiate B ed in humans. (c) Th ed in humans. bitter “Disliking” “Liking” A sweetness in rodents and humans. (a) Facial “liking” and “disliking” expressi “disliking” and “liking” (a) Facial humans. and in rodents has been identifi causation d) Pleasure (b, “liking increase may activation neural where pallidum, ventral and (Also see Plate 4.) Hedonic brain circuitry. Th circuitry. brain 4.) Hedonic fig.2.2 (Also see Plate networks have also been have identifi networks (d) Th salience “wanting.” or incentive only perhaps but pleasure diff which cortex, the orbitofrontal of food. PAG, ; VTA, ventral tegmental area. area. tegmental ventral periaqueductal gray; VTA, food. PAG,

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the neurobiology of pleasure and happiness 

1 dopamine neurotransmission (though other neurotransmitters and structures also are 2 involved). 3 Importantly, incentive salience is not hedonic impact or pleasure “liking” (Berridge 4 2 0 0 7 ) . Th is is why an individual can “want” a reward without necessarily “liking” the same 5 reward. Irrational “wanting” without liking can occur especially in addiction via incentive- 6 sensitization of the mesolimbic dopamine system and connected structures (Robinson and 7 Berridge 2003 ). At extreme, the addict may come to “want” what is neither “liked” nor 8 expected to be liked, a dissociation possible because “wanting” mechanisms are largely 9 subcortical and separable from cortically-mediated declarative expectation and conscious 10 planning. Th is is a reason why addicts may compulsively “want” to take drugs even if, at a 11 more cognitive and conscious level, they do not want to do so. Th at is surely a recipe for 12 great unhappiness (Figure 2.2 , bottom right).

13 C o r t i c a l p l e a s u r e

14 In cortex, hedonic evaluation of pleasure valence is anatomically distinguishable from 15 precursor operations such as sensory computations, suggesting the existence of a hedonic 16 cortex proper (Figure 2.2 ) (Kringelbach 2004b ). Hedonic cortex involves regions such as the 17 orbitofrontal (Kringelbach 2005 ), insula (Craig 2002 ), medial prefrontal (Amodio and Frith 18 2006 ), and cingulate cortices (Beckmann et al. 2009 ), which a wealth of human neuroimag- 19 ing studies have shown to code for hedonic evaluations (including anticipation, appraisal, 20 experience, and memory of pleasurable stimuli) and have close anatomical links to subcor- 21 tical hedonic hotspots. It is important, however, to again make a distinction between brain 22 activity coding and causing pleasure. Neural coding is inferred in practice by measuring brain 23 activity correlated to a pleasant stimulus, using human techniques (Gottfried 24 2010 ), or electrophysiological or neurochemical activation measures in animals (Aldridge 25 and Berridge 2010 ). Causation is generally inferred on the basis of a change in pleasure as a 26 consequence of a brain manipulation such as a lesion or stimulation (Green et al. 2010 ; Smith 27 et al. 2010 ). Coding and causation oft en go together for the same substrate, but they may 28 diverge so that coding occurs alone. 29 Pleasure encoding may reach an apex of cortical localization in a mid-anterior subregion 30 within the orbitofrontal cortex, where neuroimaging activity correlates strongly to subjec- 31 tive pleasantness ratings of food varieties (Kringelbach et al. 2003 ) — and to other pleasures 32 such as sexual (Georgiadis et al. 2006 ), drugs (Völlm et al. 2004 ), chocolate (Small 33 et al. 2001 ), and (Blood and Zatorre 2001 ). Most importantly, mid-anterior orbitof- 34 rontal activity tracks changes in subjective pleasure, such as a decline in palatability when 35 the reward value of one food was reduced by eating it to satiety (while remaining high to 36 another food) (Kringelbach 2005 ) , (Kringelbach et al. 2 0 0 3 ) . Th e mid-anterior subregion 37 of orbitofrontal cortex is thus a prime candidate for the coding of subjective experience of 38 pleasure (Kringelbach 2005 ). 39 Another coding site for positive hedonics in orbitofrontal cortex is along its medial edge 40 that has activity related to the positive and negative valence of aff ective events (Kringelbach 41 and Rolls 2004 ), contrasted to lateral portions that have been suggested to code unpleasant 42 events (O’Doherty et al. 2001 ) (although lateral activity may refl ect a signal to escape the 43 situation, rather than displeasure per se (Iversen and Mishkin 1970 ; Kringelbach and

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 morten l. kringelbach and kent c. berridge

1 Rolls 2003 ; Hornak et al. 2004 ; Kringelbach and Rolls 2004 )). Th is medial–lateral hedonic 2 gradient interacts with an abstraction–concreteness gradient in the posterior–anterior 3 dimension, so that more complex or abstract reinforcers (such as monetary gain and loss) 4 (O’Doherty et al. 2001 ) are represented more anteriorly in the orbitofrontal cortex than less 5 complex sensory rewards (such as taste) (Small et al. 2 0 0 1 ) . Th e medial region that codes 6 pleasant sensations does not, however, appear to change its activity with reinforcer devalua- 7 tion, and so may not refl ect the full dynamics of pleasure. 8 Other cortical regions implicated in coding for pleasant stimuli include parts of the 9 mid-insular (Craig 2009 ) and anterior cingulate cortices (Beckmann et al. 2009 ). As yet, 10 however, it is not as clear as for the orbitofrontal cortex whether those regions specifi cally 11 code pleasure or only emotion more generally (Feldman Barrett and Wager 2006). A related 12 suggestion has emerged that the frontal left hemisphere plays a special lateralized role in 13 positive aff ect more than the right hemisphere (Davidson and Irwin 1999 ), though how to 14 reconcile left -positive fi ndings with many other fi ndings of bilateral activations of orbitof- 15 rontal and related cortical regions during hedonic processing remains an ongoing puzzle 16 (Kringelbach 2005 ). 17 It remains still unknown, however, if mid-anterior orbitofrontal cortex or medial orbitof- 18 rontal cortex or any other cortical region actually causes a positive pleasure state. Clearly, 19 damage to orbitofrontal cortex does impair pleasure-related decisions, including choices 20 and context-related cognitions in humans, monkeys, and rats (Butter et al. 1 9 6 3 ; N a u t a 1 9 7 1 ; 21 Baylis and Gaff an 1991 ; Anderson et al. 1999 ; Baxter et al. 2000 ; Beer et al. 2 0 0 3 ; H o r n a k 22 et al. 2 0 0 3 ; P i c k e n s et al. 2003 , 2005 ). But some caution regarding whether cortex generates 23 positive aff ect states per se is indicated by the consideration that patients with lesions to the 24 orbitofrontal cortex do still react normally to many pleasures, although sometimes showing 25 inappropriate emotions (Damasio 1996 ; Anderson et al. 1999 ; Beer et al. 2 0 0 3 ; H o r n a k et al. 26 2003 ). Hedonic capacity aft er prefrontal damage has not, however, yet been studied in 27 careful enough detail (e.g. using selective satiation paradigms (Kringelbach et al. 2003 )), and 28 it would be useful to have more information on the role of orbitofrontal cortex, insular 29 cortex, and cingulate cortex in generating and modulating hedonic states. 30 Pleasure causation has been so far rather diffi cult to assess in humans given the limits 31 of information from lesion studies, and the correlative nature of neuroimaging studies. 32 A promising tool, however, is (DBS) which is a versatile and reversible 33 technique that directly alters brain activity in a brain target and where the ensuing whole- 34 brain activity can be measured with magnetoencephalography (MEG) (Kringelbach et al. 35 2007b ). Pertinent to a view of happiness as freedom from distress, at least pain relief can be 36 obtained from DBS of periaqueductal gray in the brainstem in humans (Gildenberg 2005 ), 37 where specifi c neural signatures of pain have been found (Green et al. 2009 ), and where the 38 pain relief is associated with activity in the mid-anterior orbitofrontal cortex, perhaps 39 involving endogenous opioid release (Kringelbach et al. 2007a ). Similarly, DBS may allevi- 40 ate some unpleasant symptoms of , though without actually producing positive 41 aff ect. 42 Famously, also, pleasure electrodes were reported to exist decades ago in animals and 43 humans when implanted in subcortical structures including the nucleus accumbens, 44 septum, and medial forebrain bundle (Olds and Milner 1954 ; Heath 1972 ) (Figure 2.2c ). 45 However, recently we and others have questioned whether most such electrodes truly caused 46 pleasure, or instead, only a psychological process more akin to “wanting” without “liking”

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the neurobiology of pleasure and happiness 

1 (Berridge and Kringelbach 2008 ). In our view, it still remains unknown whether DBS causes 2 true pleasure, or if so, where in the brain electrodes produce it.

3 Loss of pleasure

4 Th e lack of pleasure, , is one of the most important symptoms of many mental 5 illnesses including depression. It is diffi cult to conceive of anyone reporting happiness or 6 well-being while so deprived of pleasure. Th us anhedonia is another potential avenue of 7 evidence for the link between pleasure and happiness (Gorwood 2008 ). 8 Th e brain regions necessary for pleasure— but disrupted in anhedonia— are not yet clear. 9 Core “liking” reactions to sensory pleasures appear relatively diffi cult to abolish absolutely 10 in animals by a single brain lesion or drug, which may be very in evolutionary terms. 11 Only the ventral pallidum has emerged among brain hedonic hotspots as a site where 12 damage fully abolishes the capacity for positive hedonic reaction in rodent studies, replac- 13 ing even “liking” for sweetness with “disliking” gapes normally reserved for bitter or simi- 14 larly noxious tastes (Cromwell and Berridge 1993 ; Aldridge and Berridge 2010 ). Interestingly, 15 there are extensive connections from the ventral pallidum to the medial orbitofrontal cortex 16 (Öngür and Price 2000 ). 17 On the basis of this evidence, the ventral pallidum might also be linked to human anhe- 18 donia. Th is brain region has not yet been directly surgically targeted by clinicians but there 19 is anecdotal evidence that some patients with pallidotomies (of nearby , just 20 above and behind the ventral pallidum) for Parkinson’s patients show fl attened aff ect (Parkin 21 et al. 2002 ) (T. Z. Aziz, personal ), and stimulation of globus pallidus inter- 22 nus may help with depression (Kosel et al. 2007 ). A case study has also reported anhedonia 23 following bilateral lesion to the ventral pallidum (Miller et al. 2 0 0 6 ) . 24 Alternatively, core “liking” for fundamental pleasures might persist intact but unacknowl- 25 edged in anhedonia, while instead only more cognitive construals, including retrospective 26 or anticipatory savoring, becomes impaired. Th at is, fundamental pleasure may not be 27 abolished in depression aft er all. Instead, what is called anhedonia might be secondary to 28 motivational defi cits and cognitive misappraisals of rewards, or to an overlay of negative 29 aff ective states. Th is may still disrupt life enjoyment, and perhaps render higher pleasures 30 impossible. 31 Other potential regions targeted by DBS to help with depression and anhedonia include 32 the nucleus accumbens (Schlaepfer et al. 2008 ) and the subgenual cingulate cortex (Mayberg 33 et al. 2005 ). In addition, lesions of the posterior part of the anterior cingulate cortex have 34 been used for the treatment of depression with some success (Steele et al. 2008 ).

35 Bridging pleasure to meaning

36 It is potentially interesting to note that all these structures either have close links with fron- 37 tal cortical structures in the hedonic network (e.g. nucleus accumbens and ventral pallidum) 38 or belong to what has been termed the brain’s default network which changes over early 39 development (Fransson et al. 2007 ; Fair et al. 2008 ) (Figure 2.3 ).

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 morten l. kringelbach and kent c. berridge late cortices. late . ). 2007 Some et al Children Confabulations Vegetative state j l k e brain’s default network (Gusnard and and (Gusnard network default e brain’s Monitoring Cingulate regions Medial prefrontal cortex i g h Depression Cingulotomies Opiate binding f e d e brain’s default network and eudaimonic–hedonic interaction. (a–c) Th interaction. eudaimonic–hedonic and network default e brain’s . 2007 ) has been linked to self-awareness, remembering the past and prospecting the future (Addis (Addis prospecting the future the past and remembering ) has been self-awareness, to linked 2007 . Present et al Future prospection Past remembrance a c b (Also see Plate 5.) Th (Also see Plate components overlap with pleasure networks, including midline structures such as the orbitofrontal, medial prefrontal, and cingu and medial prefrontal, as the orbitofrontal, such midline structures including networks, pleasure with overlap components fig. 2.3 Raichle 2001 ; Addis

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the neurobiology of pleasure and happiness 

fig. 2.3 (continued) We whether happiness might include a role for the default 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 aff ect. 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 (Willoch et al . 2004 ), (e) have been linked to depression (Drevets et al . 1997 ), and (f) its surgical treat- ment (Steele et al. 2008 ). (g) Subregional localization of function may be indicated by con- nectivity analyses of cingulate cortex (Beckmann et al. 2009 ) and related structures, (h) important in pleasure-related monitoring, learning, and memory (Kringelbach and Rolls 2004 ), (i) as well as self-knowledge, person perception, and other cognitive functions (Amodio and Frith 2006 ). (j) Th e default network may change over early life in children and pre-term babies (Fransson et al . 2007 ; Fair et al. 2008 ), (k) in pathological states includ- ing depression and vegetative states (Laureys et al . 2004 ), (l) and aft er lesions to its medial orbitofrontal and subgenual cingulate cortices that disrupt reality monitoring and create spontaneous confabulations (Schnider 2003 ).

1 Mention of the default network brings us back to the topic of eudaimonic happiness, and 2 to potential interactions of hedonic brain circuits with circuits that assess meaningful 3 relationships of self to social others. Th e default network is a steady-state circuit of the brain 4 which becomes perturbed during cognitive tasks (Gusnard and Raichle 2001 ). Most perti- 5 nent here is an emerging that has proposed the default network to carry represen- 6 tations of self (Lou et al. 1999 ), internal modes of cognition (Buckner et al. 2008 ), and 7 perhaps even states of consciousness (Laureys et al. 2004 ). Such functions might well be 8 important to higher pleasures as well as meaningful aspects of happiness. 9 Although highly speculative, we wonder whether the default network might deserve 10 further consideration for a role in connecting eudaimonic and hedonic happiness. At least, 11 key regions of the frontal default network overlap with the hedonic network discussed earlier, 12 such as the anterior cingulate and orbitofrontal cortices (Kringelbach and Rolls 2004 ; Amodio 13 and Frith 2006 ; Steele et al. 2008 ; Beckmann et al. 2009 ), and have a relatively high density of 14 opiate receptors (Willoch et al. 2004 ). And activity changes in the frontal default network, 15 such as in the subgenual cingulate and orbitofrontal cortices, correlate to pathological changes 16 in subjective hedonic experience, such as in depressed patients (Drevets et al. 1 9 9 7 ) . 17 Pathological self-representations by the frontal default network could also provide a 18 potential link between hedonic distortions of happiness that are accompanied by eudaimo- 19 nic dissatisfaction, such as in cognitive rumination of depression (Williams et al. 1996 ; 20 Schnider 2003 ; Addis et al. 2007 ). Conversely, mindfulness-based cognitive therapy for 21 depression, which aims to disengage from dysphoria-activated depressogenic thinking, 22 might conceivably recruit default network circuitry to help mediate improvement in 23 happiness via a linkage to hedonic circuitry (Teasdale et al . 2000 ).

24 Concluding remarks

25 Th e most diffi cult questions facing pleasure and happiness research remain the nature of its 26 subjective experience and the relation of hedonic components (pleasure or positive aff ect)

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 morten l. kringelbach and kent c. berridge

1 to eudaimonic components (cognitive appraisals of meaning and life satisfaction). While 2 some progress has been made in understanding brain hedonics, it is important not to over- 3 interpret. In particular we have still not made substantial progress towards understanding 4 the functional neuroanatomy of happiness. 5 In this review we have, however, identifi ed a number of brain regions that are important 6 in the brain’s hedonic networks, and speculated on potential interaction with eudaimonic 7 networks. While it remains unclear how pleasure and happiness are exactly linked, it may be 8 safe to say at least that the pathological lack of pleasure, in anhedonia or dysphoria, amounts 9 to a formidable obstacle to happiness. 10 In social animals like humans, social interactions with conspecifi cs are fundamental and 11 central to enhancing the other pleasures. Humans are intensely social, and data indicate that 12 one of the most important factors for happiness is social relationships with other people. 13 Social pleasures may still include vital sensory features such as visual faces, touch features of 14 grooming and caress, as well as in humans more abstract and cognitive features of social 15 reward and relationship evaluation (Adolphs 2003 ). 16 In particular, adult pair bonds and attachment bonds between parents and infants are 17 likely to be extremely important for the survival of the species (Kringelbach et al. 2008 ). Th e 18 breakdown of these bonds is all too common and can lead to great unhappiness. And even 19 bond formation can potentially disrupt happiness, such as in transient parental depression 20 aft er birth of an infant (in over 10 of mothers and approximately 3 of fathers (Cooper and 21 Murray 1998 )). Progress in understanding the hedonics of social bonds could be useful in 22 understanding happiness. 23 is beginning to unravel some of the complex dynamics of human 24 social interactions. One of its major challenges is to map the developmental changes in 25 reward processing over a lifespan. Another challenge is to understand how brain networks 26 underlying fundamental pleasure relate to higher pleasures such as music, , play and 27 fl ow, and to happiness. 28 Further, so far as positive aff ect contributes to happiness, then considerable progress has 29 been made in understanding the neurobiology of pleasure in ways that might be relevant. 30 For example, we can imagine several possibilities to relate happiness to particular hedonic 31 psychological processes discussed previously. Th us, one way to conceive of hedonic happi- 32 ness is as “liking” without “wanting.” Th at is, a state of pleasure without disruptive , a 33 state of (Kringelbach 2009 ). Another possibility is that moderate “wanting” 34 matched to positive “liking” facilitates engagement with the world. A little incentive salience 35 may add zest to the perception of life and perhaps even promote the construction of mean- 36 ing, just as in some patients DBS may help lift the veil of depression by making life events 37 more appealing. However, too much “wanting” can readily spiral into maladaptive patterns 38 such as addiction, and is a direct route to great unhappiness. Finally, happiness might spring 39 from higher pleasures, positive appraisals of life meaning, and social connectedness, all 40 combined and merged by interaction between the brain’s default networks and pleasure 41 networks. Achieving the right hedonic balance in such ways may be crucial to keep one not 42 just ticking over but perhaps even happy. 43 Future scientifi c advances may provide a better sorting of psychological features of happi- 44 ness and its underlying brain networks. If so, it remains a distinct possibility that more 45 among us may be one day shift ed into a better situation to enjoy daily events, to fi nd life 46 meaningful and worth living— and perhaps even to achieve a degree of bliss.

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the neurobiology of pleasure and happiness 

1 Acknowledgments

2 We thank Christopher Peterson, Eric Jackson, Kristine R ø mer Th omsen, Christine Parsons, 3 and Katie Young for helpful comments on an earlier version of this manuscript. Our research 4 has been supported by grants from the TrygFonden Charitable Foundation to MLK and 5 from the NIMH and NIDA to KCB.

6 References

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 morten l. kringelbach and kent c. berridge

1 Steiner , J.E. , Glaser , D. , Hawilo , M.E. , and Berridge , K.C. ( 2 0 0 1 ) . C o m p a r a t i v e e x p r e s s i o n o f 2 hedonic impact: aff ective reactions to taste by human infants and other primates . 3 Neuroscience and Biobehavioral Reviews , 25 , 53 – 74 . 4 Teasdale , J.D. , Segal , Z.V. , Williams , J.M. , Ridgeway , V.A. , Soulsby , J.M. , and Lau , M.A. 5 ( 2000 ). Prevention of relapse/recurrence in major depression by mindfulness-based cogni- 6 tive therapy. Journal of Consulting and Clinical Psychology , 68 , 615 – 23 . 7 Veldhuizen , M.G. , Rudenga , K.J. , and Small , D.M. ( 2 0 1 0 ) . Th e pleasure of taste, fl avor 8 and food . In M.L. Kringelbach and K.C. Berridge (eds.) Pleasures of the Brain , pp. 146 – 68 . 9 New York: Oxford University Press. 10 Völlm , B.A . , de Araujo , I.E.T. , Cowen , P. J. , et al . ( 2004 ). Methamphetamine activates reward 11 circuitry in drug naïve human subjects . Neuropsychopharmacology , 29 , 1715 – 22 . 12 Vuust , P. and Kringelbach , M.L. ( 2 0 1 0 ) . Th e pleasure of music. In M.L. Kringelbach , and 13 K.C. Berridge (eds.) Pleasures of the Brain, pp. 255– 69 . New York: Oxford University Press. 14 Waterman , A.S. ( 1993 ). Two conceptions of happiness: contrasts of personal expressiveness 15 (eudaimonia) and hedonic enjoyment. Journal of Personality and Social Psychology , 64 , 16 678 – 91 . 17 Watson , K.K. , Shepherd , S.V. , and Platt , M.L. (2010 ). of pleasure. In 18 M.L. Kringelbach , and K.C. Berridge (eds.) Pleasures of the Brain , pp. 85 – 95 . New York : 19 Oxford University Press. 20 Williams , J.M. , Ellis , N.C. , Tyers , C. , Healy , H. , Rose , G. , and MacLeod , A.K. ( 1996 ). Th e 21 specifi city of autobiographical memory and imageability of the future. Memory & Cognition , 22 24 , 116 – 25 . 23 Willoch , F. , Schindler , F. , Wester , H.J. , et al . ( 2004 ). Central poststroke pain and reduced 24 opioid receptor binding within pain processing circuitries: a [11C]diprenorphine PET study. 25 Pain , 108 , 2 1 3 – 2 0 .

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