Well-being and affective style: neural substrates and biobehavioural correlatesIntroductionConceptual and methodological considerations in the study of affective styleNeural substrates of emotion and affective stylePrefrontal cortexAmygdalaHippocampus and ACCWhat are individual differences in PFC and amygdala activations associated with?Emotion regulation: a key component of affective stylePlasticity in the central circuitry of emotionSummary and conclusionsREFERENCESPublished online 11 August 2004Well-being and affective style: neural substratesand biobehavioural correlatesRichard J. DavidsonLaboratory for Affective Neuroscience, W. M. Keck Laboratory for Functional Brain Imaging and Behavior,University of Wisconsin-Madison, 1202 West Johnson Street, Madison, WI 53706, USA ([email protected])One of the most salient features of emotion is the pronounced variability among individuals in their reactionsto emotional incentives and in their dispositional mood. Collectively, these individual differences have beendescribed as affective style. Recent research has begun to dissect the constituents of affective style. Thesearch for these components is guided by the neural systems that instantiate emotion and emotion regu-lation. In this article, this body of research and theory is applied specifically to positive affect and well-being.The central substrates and peripheral biological correlates of well-being are described. A resilient affectivestyle is associated with high levels of left prefrontal activation, effective modulation of activation in the amyg-dala and fast recovery in response to negative and stressful events. In peripheral biology, these central pat-terns are associated with lower levels of basal cortisol and with higher levels of antibody titres to influenzavaccine. The article concludes with a consideration of whether these patterns of central and peripheralbiology can be modified by training and shifted toward a more salubrious direction.Keywords: affective neuroscience; resilience; prefrontal cortex; brain asymmetry; emotion regulation;affective style1. INTRODUCTIONOne of the most salient characteristics of emotion is theextraordinary heterogeneity in how different individualsrespond to the same emotionally provocative challenge.Such differences in patterns of emotional reactivity play acrucial role in shaping variations in well-being. Althoughindividual differences in emotion processing can be foundat many levels of phylogeny, they are particularly pro-nounced in primates and probably are most extreme inhumans. A number of evolutionary theorists have specu-lated on the adaptive significance of such individual differ-ences (Wilson 1994). Although these arguments havenever been applied to the domain of emotion and affectivestyle, it is not difficult to develop hypotheses about howsuch differences might provide advantages to individualsliving in groups. However, rather than focus on the distalcauses of such individual differences which are so difficultto subject to rigorous test, I wish only to call attention tothe possibility that variability in characteristics such as‘fearfulness’ or ‘cheerfulness’ might provide some adaptivebenefit to individuals living together in groups. Instead,this article examines the proximal mechanisms that under-lie such individual differences, with a focus on well-being.The central substrates of individual differences in compo-nents of well-being will be described. The possible influ-ence of the central circuitry of emotion on peripheralbiological indices that are relevant to physical health andillness will also be considered. It is helpful to contrast well-being with specific types of psychopathology that involvedysfunctions in the circuitry of adaptive emotionalresponding. Accordingly, some mention of recent work onthe neurobiology of mood and anxiety disorders will bemade. Finally, plasticity in the underlying brain circuitrythat instantiates affective style will be described and its rolein promoting resilience will be considered.Affective style refers to consistent individual differencesin emotional reactivity and regulation (see Davidson1998a; Davidson et al. 2000 a ,b). It is a phrase that is meantto capture a broad array of processes that, either singly orin combination, modulate an individual’s response toemotional challenges, dispositional mood and affect-relevant cognitive processes. Affective style can refer tovalence-specific features of emotional reactivity or mood,or it can refer to discrete emotion-specific features. Bothlevels of analysis are equally valid and the choice of levelshould be dictated by the question posed.Rapid developments in our understanding of emotion,mood and affective style have come from the study of theneural substrates of these phenomena. The identificationof the brain circuitry responsible for different aspects ofaffective processing has helped to parse the domain of emo-tion into more elementary constituents in a manner similarto that found in cognitive neuroscience, where an appeal tothe brain has facilitated the rapid development of theoryand data on the subcomponents of various cognitive pro-cesses (e.g. Kosslyn & Koenig 1992).This article will highlight some of the advances that havebeen made in our understanding of the brain mechanismsthat underlie affective style. These advances have emergedfrom three major sources: studies of patients with discretelesions of the brain; neuroimaging studies of normal indivi-One contribution of 12 to a Discussion Meeting Issue ‘The science ofwell-being: integrating neurobiology, psychology and social science’.Phil. Trans. R. Soc. Lond. B (2004) 359, 1395–1411 1395#2004 The Royal Societydoi:10.1098/rstb.2004.1510duals; and studies of pathologies of brain function inpatients with various psychiatric and neurological disordersthat involve abnormalities in emotion. I will use thematerial on pathology to help to identify the neural cir-cuitry crucial to certain forms of positive affect so that wecan begin to place well-being squarely within a neurobiolo-gical framework.Both lesion and neuroimaging studies provide infor-mation primarily on the ‘where’ question; that is, where inthe brain are computations related to specific aspects ofaffective processing occurring. It is important at the outsetto consider both the utility of knowing ‘where’ and howsuch information can provide insight into the ‘how’ ques-tion; that is, how might a particular