Emotion
Positive Affect and Markers of Inflammation: Discrete
Positive Emotions Predict Lower Levels of Inflammatory
Cytokines
Jennifer E. Stellar, Neha John-Henderson, Craig L. Anderson, Amie M. Gordon, Galen D.
McNeil, and Dacher Keltner
Online First Publication, January 19, 2015. http://dx.doi.org/10.1037/emo0000033

CITATION
Stellar, J. E., John-Henderson, N., Anderson, C. L., Gordon, A. M., McNeil, G. D., & Keltner, D.
(2015, January 19). Positive Affect and Markers of Inflammation: Discrete Positive Emotions
Predict Lower Levels of Inflammatory Cytokines. Emotion. Advance online publication.
http://dx.doi.org/10.1037/emo0000033

 Emotion
2015, Vol. 14, No. 6, 000

© 2015 American Psychological Association
1528-3542/15/$12.00 http://dx.doi.org/10.1037/emo0000033

BRIEF REPORT

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This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

Positive Affect and Markers of Inflammation: Discrete Positive Emotions
Predict Lower Levels of Inflammatory Cytokines
Jennifer E. Stellar

Neha John-Henderson

University of Toronto

University of Pittsburgh

Craig L. Anderson, Amie M. Gordon, Galen D. McNeil, and Dacher Keltner
University of California–Berkeley
Negative emotions are reliably associated with poorer health (e.g., Kiecolt-Glaser, McGuire, Robles, &
Glaser, 2002), but only recently has research begun to acknowledge the important role of positive
emotions for our physical health (Fredrickson, 2003). We examine the link between dispositional positive
affect and one potential biological pathway between positive emotions and health—proinflammatory
cytokines, specifically levels of interleukin-6 (IL-6). We hypothesized that greater trait positive affect
would be associated with lower levels of IL-6 in a healthy sample. We found support for this hypothesis
across two studies. We also explored the relationship between discrete positive emotions and IL-6 levels,
finding that awe, measured in two different ways, was the strongest predictor of lower levels of
proinflammatory cytokines. These effects held when controlling for relevant personality and health
variables. This work suggests a potential biological pathway between positive emotions and health
through proinflammatory cytokines.
Keywords: proinflammatory cytokines, positive emotion, health

cytokines have been implicated in the onset and progression of
numerous chronic diseases including diabetes, cardiovascular disease, and depression (Cesari et al., 2003; Dowlati et al., 2010;
Wellen & Hotamisligil, 2005). Although past research has focused
on the relationship between proinflammatory cytokines and negative emotions, little is known about whether proinflammatory
cytokines may be associated with positive emotions. We hypothesize that trait positive affect will be associated with lower levels
of proinflammatory cytokines and explore whether certain discrete
positive emotions are particularly predictive of reduced levels of
these promoters of inflammation.
Stressful experiences and certain negative emotions, moods, and
traits have been linked to higher systemic levels of proinflammatory cytokines (e.g., Kiecolt-Glaser et al., 2002). For example,
self-reported fear elicited during the Trier Social Stress Task
predicts elevated proinflammatory cytokines (Moons et al., 2010).
Experiences of shame induced by writing about instances of selfblame on three separate days led to increases in proinflammatory
cytokines pre- to posttask (Dickerson, Kemeny, Aziz, Kim, &
Fahey, 2004). Negative affect is also associated with proinflammatory cytokines in studies using trait measures, although this
relationship is likely bidirectional (Messay, Lim, & Marsland,
2012). Clinical depression and anxiety are associated with higher
levels of proinflammatory cytokines (Dowlati et al., 2010;
O’Donovan et al., 2010), as is greater self-reported trait negative
affect (Denollet, Vrints, & Conraads, 2008).
Although little is known about whether positive emotions are
associated with levels of proinflammatory cytokines, there is rea-

Negative affect reliably predicts poorer health, including risk of
mortality, heart disease, and cancer (see Kiecolt-Glaser, McGuire,
Robles, & Glaser, 2002 for a review). One pathway by which
negative emotions lead to poor health outcomes is through increasing inflammation, a response of the immune system (Moons,
Eisenberger, & Taylor, 2010). Communication molecules called
proinflammatory cytokines are essential in the coordination and
promotion of inflammatory processes. Acute inflammatory activity
is essential in responding adaptively to infection and injury. However, chronically elevated levels of proinflammatory cytokines in
the absence of illness or injury can lead to negative health outcomes by stimulating the hypothalamic–pituitary–adrenal axis,
increasing levels of C-reactive protein, and slowing muscle repair
(Kiecolt-Glaser et al., 2002). Elevated levels of proinflammatory

Jennifer E. Stellar, Department of Management, University of Toronto;
Neha John-Henderson, Department of Cardiovascular Behavioral Medicine, University of Pittsburgh; Craig L. Anderson, Amie M. Gordon, Galen
D. McNeil, and Dacher Keltner, Department of Psychology, University of
California–Berkeley.
This research was supported by John Templeton Foundation grants
(95524 and 88210).
Correspondence concerning this article should be addressed to Jennifer E. Stellar, 105 St. George St., Rotman School of Management,
University of Toronto, Toronto ON M5S 3E6, Canada. E-mail:
jennifer.stellar@utoronto.ca
1

 STELLAR ET AL.

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2

son to believe that the two may have an inverse relationship. More
generally, the experience of positive emotions leads to greater
longevity and reduced morbidity (e.g., Pressman & Cohen, 2005).
The capacity for positive emotions to promote faster recovery from
stress (Folkman & Moskowitz, 2000) and help strengthen social
ties (Fredrickson, 2003; Lyubomirsky, King, & Diener, 2005)
point to possible mechanisms by which positive emotions lead to
better health and reduced inflammatory activity.
The existing literature offers suggestive evidence that positive
affect is associated with lower levels of proinflammatory cytokines. For instance, optimism (typically associated with greater
positive affect) predicts lower levels of proinflammatory cytokine
responses to a stressor (Brydon, Walker, Wawrzyniak, Chart, &
Steptoe, 2009), and watching an amusing film temporarily decreases levels of proinflammatory cytokines (Mittwoch-Jaffe,
Shalit, Srendi, & Yehuda, 1995). Moreover, women who frequently reported being very or extremely happy over the course of
a day had lower levels of proinflammatory cytokines (Steptoe,
O’Donnell, Badrick, Kumari, & Marmot, 2008).
Although these studies suggest an association between trait
positive affect and reduced levels of proinflammatory cytokines,
no research has examined the varying degree to which different
positive emotions are associated with these promoters of inflammation. A discrete approach to emotions has proven fruitful in
identifying specific negative emotions, such as shame and fear,
which are particularly potent elicitors of proinflammatory cytokines (e.g., Dickerson et al., 2004). Assuming that trait positive
affect is associated with reduced levels of proinflammatory cytokines, examining individual positive emotions may help identify
emotion-specific mechanisms that help explain this relationship.
In this investigation, we present two studies that test the hypothesis that trait positive affect will be associated with reduced
levels of inflammatory cytokines. In Study 2, we examine this
relationship in more depth by exploring whether certain discrete
positive emotions are stronger predictors of levels of proinflammatory cytokines.

Samples were frozen and stored at ⫺80 °C in a freezer in the
laboratory. Levels of IL-6 were determined by an enzyme-linked
immunosorbent assay (ELISA) using commercially available kits
(R&D Systems, Minneapolis, MN). Samples from each individual
were run in duplicate on the same plate. The intra-assay coefficient
of variation (CV) was 8.5%, and the interassay CV was 9.6%.
After the OMT collection, participants completed an activity for
another experiment, were debriefed, and were released.

Measures
Body mass index. Participants provided their height and
weight to calculate body mass index (BMI; M ⫽ 22.72, SD ⫽
3.65). We included BMI as a covariate because past work has
shown that it correlates positively with levels of interleukin-6
(IL-6) and weight varies greatly in the undergraduate population
(Khaodhiar, Ling, Blackburn, & Bistrian, 2004).
Positive and Negative Affect Schedule. Participants reported
how much they had experienced negative and positive affect over
the course of the past month using 20 items (e.g., inspired, enthusiastic, hostile) on the Positive and Negative Affect Schedule
(PANAS) ranging from 1 (never) to 5 (always; Watson, Clark, &
Tellegen, 1988). The positive (M ⫽ 3.46, SD ⫽ 0.68) and negative
(M ⫽ 2.51, SD ⫽ 0.81) subscales showed strong reliability (␣s ⱖ
0.87) and were negatively correlated, r(94) ⫽ ⫺0.23, p ⫽ .03.

Results
The distribution of IL-6 was not normal; therefore, the values
were log transformed in line with prior research (John-Henderson,
Rheinschmidt, Mendoza-Denton, & Francis, 2014). In keeping
with our hypotheses, a regression revealed that positive affect
negatively predicted IL-6, controlling for BMI, ␤ ⫽ ⫺.30, p ⫽
.004.1 In a second regression analysis, negative affect was not a
significant predictor of IL-6, controlling for BMI, ␤ ⫽ .06, p ⫽
.71. These results demonstrate that trait positive affect predicts
lower levels of proinflammatory cytokines in a healthy sample.

Study 1

Study 2

Participants

Participants

Ninety-four freshman undergraduates (30 males, 62 females, 2
declined to state) from a large West Coast university participated
in this study for credit in a psychology class. The sample was 4%
African American, 46% Asian American, 42% European American, 13% Latin American, and 8% other ethnicities.

One hundred nineteen freshman undergraduates (36 males, 83
females) from a large West Coast university participated in this
study for payment. The sample was 5% African American, 65%
Asian American, 22% European American, 14% Latin American,
and 4% other ethnicities.

Procedure

Procedure

Participants arrived at the laboratory and completed background
questionnaires on the computer. They provided a sample of oral
mucosal transudate (OMT) approximately 15 min after they arrived at the laboratory. OMT is a filtrate of blood plasma that
comes from the capillaries at the base of the crevice in between the
teeth and gums, which then passes across the mucosa and gingival
crevices and settles in the mouth. Participants were instructed to
refrain from eating or drinking 1 hr before providing samples. An
Orasure collective device (Epitope, Beaverton, OR) was placed
between the lower cheek and gum for 2 min on the left side.

Participants took part in this study as part of a multisession
longitudinal study. Participants completed background questionnaires on their home computers using a secure website. In a
1
When including additional health-related covariates to predict IL-6
levels (whether participants had a chronic health condition, whether they
smoked, and average hours of sleep), the overall pattern of results was the
same. Thirteen of fourteen regression analyses remained significant,
␤s ⱖ ⫺0.20, ps ⱕ .05; in Study 2, the PANAS positive affect became a
marginally significant predictor, ␤ ⫽ ⫺0.20, p ⬍ .06.

 POSITIVE AFFECT AND INFLAMMATORY CYTOKINES

follow-up session, participants came to the laboratory for a collection of OMT, which was conducted and assayed in the same
manner as in Study 1.

Table 1
Positive Emotions Predicting IL-6
DPES Subscale

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Measures
BMI. BMI (M ⫽ 22.5, SD ⫽ 3.7) was collected for the same
reason as Study 1.
PANAS. We used a validated, 10-item shortened version of
the PANAS that assesses general positive and negative affect in
the last month (Thompson, 2007). The positive (M ⫽ 3.75, SD ⫽
0.57) and negative (M ⫽ 2.26, SD ⫽ 0.66) subscales showed
strong reliability (␣s ⱖ 0.72) and were not correlated, r(119) ⫽ ⫺.04,
p ⫽ .69.
Dispositional Positive Emotions. The Dispositional Positive
Emotions Scale (DPES) assesses the extent to which participants
experience key positive emotions in their daily lives (Shiota,
Keltner, & John, 2006). This scale focuses on seven positive
emotions—amusement, awe, compassion, contentment, joy, love,
and pride—with five to six items for each emotion. Participants
marked their agreement on a 7-point Likert scale ranging from 1
(strongly disagree) to 7 (strongly agree) to statements such as “I
feel wonder almost every day” (awe) and “I am an intensely
cheerful person” (joy). The entire DPES scale (M ⫽ 4.98, SD ⫽
0.71) and its seven subscales showed strong reliability (␣s ⱖ 0.81).
The Big Five Personality Inventory. The Big Five Personality Inventory (BFI; M ⫽ 3.74, SD ⫽ 0.49) measures five facets
of personality, including agreeableness, conscientiousness, neuroticism, extraversion, and openness, using 44 items (John & Srivastava, 1999). Participants reported how much they agreed or
disagreed from 1 (strongly disagree) to 5 (strongly agree) to 8 –10
self-descriptive statements such as “is original.” All subscales
showed strong reliability (␣s ⱖ 0.76).

Results
Twelve participants did not complete both the online and laboratory session of the study, and two participants had IL-6 samples
that did not produce viable results, which left a sample of 105. We
log transformed the non-normal distribution of IL-6. In keeping
with Study 1, positive affect, measured through the PANAS,
negatively predicted levels of IL-6 controlling for BMI,
␤ ⫽ ⫺0.21, p ⫽ .04, whereas negative affect did not, ␤ ⫽ ⫺0.02,
p ⫽ .88. In addition, our second measure of trait positive affect, the
DPES, also negatively predicted IL-6 levels, controlling for BMI,
␤ ⫽ ⫺0.25, p ⫽ .01. These results suggest that greater trait
positive affect predicts lower levels of proinflammatory cytokines.
To explore whether certain discrete positive emotions were
more strongly associated with reduced levels of proinflammatory
cytokines, we examined each of the seven subscales of the DPES
in separate regressions controlling for BMI. Awe, joy, contentment, and pride negatively predicted IL-6 (see Table 1, column 1).
Dispositional awe had the strongest relationship with IL-6 of any
positive emotion. When applying a Bonferroni family-wise correction for multiple tests, only awe remained a significant predictor
(significance threshold of p ⫽ .007). In addition, when all seven
subscales of the DPES were simultaneously entered into a regression, only awe significantly predicted levels of IL-6 (see Table 1,
column 2).

3

Awe
Amusement
Compassion
Contentment
Joy
Love
Pride

IL-6
ⴱⴱⴱ

⫺0.33
⫺0.02
⫺0.09
⫺0.20ⴱ
⫺0.23ⴱ
⫺0.10
⫺0.21ⴱ

IL-6
⫺0.33ⴱⴱ
0.16
0.05
0.04
⫺0.11
⫺0.07
⫺0.009

Note. ␤ values for positive emotions predicting IL-6 and controlling for
participant’s BMI. In column 1, emotions are separately entered into
regressions and in column 2 they are simultaneously entered.
ⴱ
p ⬍ .05. ⴱⴱ p ⬍ .01. ⴱⴱⴱ p ⬍ .001.

To further explore this strong relationship between awe and
proinflammatory cytokine levels, we examined a second measure
of awe—participants’ reports of experiencing awe the day they
visited the laboratory. In the laboratory session, participants were
asked how much they felt awe, wonder, and amazement that day,
ranging from 1 (not at all) to 10 (very much). These three measures
were combined into a composite of the daily experience of awe
and showed a high reliability (M ⫽ 6.35, SD ⫽ 1.58; ␣ ⫽ .88). The
more frequently participants reported feeling awe, wonder, and
amazement that day, the lower their levels of IL-6, controlling for
BMI, ␤ ⫽ ⫺0.48, p ⬍ .001.
Awe has been linked to the personality facet of openness and
was correlated with openness in this study, r(119) ⫽ 0.50, p ⬍ .01
(Shiota et al., 2006). When controlling for openness to experience,
dispositional awe and the amount of awe experienced that day
continued to predict IL-6, ␤ ⫽ ⫺0.33, p ⫽ .004 and ␤ ⫽ ⫺0.47,
p ⬍ .001, respectively.

General Discussion
Across two studies, trait positive affect predicted lower levels of
the proinflammatory cytokine IL-6. Dispositional joy, contentment, pride, and awe each negatively predicted levels of IL-6.
However, awe was the strongest predictor. This finding was replicated in a second analysis, in which the amount of awe experienced that day also predicted levels of IL-6.
Although our measure assessed affect over the past month to
predict current levels of IL-6, our work cannot directly address
questions of causality. Past research has shown that infecting
individuals with influenza, which increases proinflammatory cytokine production, has led to reduced positive affect the next day
(Janicki-Deverts, Cohen, Doyle, Turner, & Treanor, 2007). Therefore, it is possible and probably likely that there is a bidirectional
relationship between positive affect and cytokine production.
Why would awe be such a potent predictor of reduced proinflammatory cytokines? One reason is that proinflammatory cytokines encourage social withdrawal and reduce exploration, which
would serve the adaptive purpose of helping an individual recover
from injury or sickness (Maier & Watkins, 1998). On the other
hand, awe is associated with curiosity and a desire to explore,
suggesting antithetical behavioral responses to those found during
inflammation (Keltner & Haidt, 2003). In this sense, experiences
of awe may be part of an integrated response that includes emo-

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4

tional and biological responses that facilitate approach and social
exploration. Alternatively, awe elicits feelings of interconnectedness with others (Keltner & Haidt, 2003) and may affect levels of
cytokines through fostering social connection and supportive relationships (Kiecolt-Glaser, Gouin, & Hantsoo, 2010). In many
ways awe lies on the opposite end of the spectrum from shame, a
known elicitor of inflammatory activity (Dickerson et al., 2004).
We did not find a strong relationship between trait negative
affect and IL-6 in either study. There are a few explanations for
this lack of an association. Past work suggests that social stressors
(i.e., social evaluation and rejection) and negative emotions that
motivate social withdrawal (e.g., shame) are particularly strong
elicitors of proinflammatory cytokine production (e.g., Dickerson
et al., 2004; Slavich, Way, Eisenberger, & Taylor, 2010). These
types of discrete emotions are not well represented in the PANAS.
In addition, much of the literature on negative affect and proinflammatory cytokines is associated with state experiences. It may
be that negative emotions exert the strongest influence on proinflammatory cytokines at the state, rather than trait, level. Much of
the research that has been conducted on trait negative affect
utilizes clinical samples, not healthy populations like our sample.
To our knowledge, no studies have yet to identify an association
between PANAS trait negative affect and proinflammatory cytokines within a healthy sample.
It is important to acknowledge that we measured levels of
proinflammatory cytokines in OMT, not in blood, which most
studies use when assessing proinflammatory cytokine levels. Research has shown that levels of proinflammatory cytokines in
OMT are related to stressors and psychosocial risk factors and
predict negative health outcomes (Dickerson et al., 2004; Saxton,
John-Henderson, Reid, & Francis, 2011; Sjögren, Leanderson,
Kristenson, & Ernerudh, 2006; Slavich et al., 2010). Given the
above associations, although not a surrogate for measurements of
systemic inflammatory activity in blood, measuring levels of proinflammatory cytokines in OMT appears to be a viable method.
The present research provides a potential biological pathway for
the association between positive emotions and health. Positive
emotions, especially awe, are associated with lower levels of
proinflammatory cytokines. This research contributes to a growing
body of work that demonstrates that positive emotions not only
feel good, they are good for the body.

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Received July 21, 2014
Revision received October 7, 2014
Accepted October 8, 2014 䡲